The significance of aquatic lateral carbon (C) export in mangrove ecosystems highlights the extensive contribution of aquatic pathways to the net ecosystem carbon budget. However, few studies have investigated lateral fluxes of dissolved organic carbon (DOC) and inorganic carbon (DIC), partly due to methodological difficulty. Therefore, we evaluated area-based lateral C fluxes in a small mangrove estuary that only had one exit for water exchange to the coast. We sampled water from the mouth of the creek and integrated discharge and consecutive concentration of mangrove-derived C (ΔC). Then, we estimated the area-normalized C fluxes based on the inundated mangrove area. DIC and DOC concentrations at the river mouth increased during ebb tide during both summer and winter. We quantified the ΔC in the estuary using a two-component conservative mixing model of freshwater and seawater. DIC and DOC proportions of ΔC concentrations at the river mouth during ebb tide was between 34% and 56% in the winter and 26% and 42% in the summer, respectively. DIC and DOC fluxes from the estuary were estimated to be 1.36 g C m−2 d−1 and 0.20 g C m−2 d−1 in the winter and 3.35 g C m−2 d−1 and 0.86 g C m−2 d−1 in the summer, respectively. Based on our method, daily fluxes are mangrove area-based DIC and DOC lateral exports that can be directly incorporated into the mangrove carbon budget.

© 2019, Coastal and Estuarine Research Federation. To clarify the effects of crab burrows on variation in sediment CO2 flux in mangrove forest, we measured the traits of crab burrows (density and entrance area size) and the CO2 flux rate from sediment surfaces, in areas with and without burrows, in a subtropical mangrove forest on Ishigaki Island, southwestern Japan. Burrow density and entrance area showed significant differences among seasons (warm, middle, and cool) and mangrove zones (upper-, middle-, and downstream), which may have depended on crab phenology, life cycle, and species composition. The sediment CO2 flux rate was significantly higher at plots with crab burrows (B+) than at those without burrows (B−) in each zone and season. However, standardized sediment CO2 flux rate by burrow surface area at B+ plots did not differ significantly from that at B− plots. In addition, there were no significant differences in sediment temperature and sediment water content between the two types of plots. Moreover, the level of microbial respiration differed significantly between sediments collected from the deep part and those collected from either the ground surface part or burrow walls. These results suggest that crab burrows increase sediment CO2 flux from the mangrove forest floor by increasing the sediment–atmosphere interface area, thereby inducing a change to aerobic conditions in the sediments around burrows. Therefore, the seasonal and spatial effect of crab burrows on the forest floor should be considered when evaluating sediment CO2 flux and examining the role of the mangrove ecosystem as a carbon sink.

© 2019 Elsevier B.V. Numerous studies have examined the variability of stemflow across different tree species under different meteorological conditions. However, studies have rarely considered stemflow associated with individual rainfall events and stemflow DOM flux. Therefore, we collected stemflow data from >100 individual rainfall events across four diameter size classes (20–30 cm, 30–40 cm, 40–50 cm, and DBH > 50 cm) of Castanopsis cuspidata in an evergreen forest (35° 26′ N, 136° 47E). The main objectives were to evaluate the stemflow hydrology and DOM flux characteristics in relation to tree size and different rainfall intensities based on a large dataset of individual rainfall events. In line with previous studies, the mean stemflow volumes and percentages of larger trees were higher than those of smaller trees; smaller trees have a higher funneling ratio than larger trees at both tree-scale (FRt) and stand-scale (FRs), which means that smaller trees are more effective in funneling water to their base. However, tree size significantly affects the stemflow volume and percentage only when the rainfall intensity is below 15 mm h−1. Tree size had a limited effect on the stemflow DOM concentration and stemflow DOM yield at a monthly scale. However, stemflow DOM flux and the flux-based DOM enrichment ratio were affected profoundly by tree size. Similar to the funneling ratio, small trees had a higher flux-based DOM enrichment ratio and supplied more DOM per unit trunk basal area. Thus, in addition to tree species, we suggest that tree size is also an important factor influencing the heterogeneity of the spatial patterns of the soil solution chemistry near the tree trunks.

© 2019, © 2019 Japanese Society of Soil Science and Plant Nutrition. The role of biochar in the mitigation of CO2 emissions has been extensively studied in agricultural soils but is not well understood in Japanese forest soils, especially in relation to CO2 emissions from applied biochar and native soil C (i.e., the priming effect; PE). We hypothesized that the type of biochar and/or the application method (mixed or sprinkled) affect the direction and magnitude of PE in forest soil, and in particular, negative PE can be achieved relatively easily if biochar produced under higher temperature conditions were sprinkled on the soil surface. To test our hypothesis, we measured CO2 emissions from biochar-amended brown forest soil in Japan and examined its PE by conducting a medium-term (~4 months) incubation study. As substrates, we used plain straw from the C4 grass Miscanthus sinensis (SU) and two qualities of biochar produced from it at either 300°C (BC300) or 800°C (BC800) and compared two application methods: mixed into or sprinkled onto the soil. BC800 had a greater C content and C:N ratio as well a lower volatile matter content and higher nonvolatile matter content than BC300. SU had the lowest C content and C:N ratio of all the substrates. We found that biochar quality (volatile and nonvolatile matter content) was clearly related to the decomposition rate when mixed into the soil, but we could not find this relationship when biochar was sprinkled onto the soil. The addition of biochar to the soil induced a positive PE in the early stages (except for BC800 sprinkled) but suppressed CO2 emissions from native soil organic matter (negative PE) in the later stages, whereas the PE with SU application was always positive regardless of application method. Our results suggest that when biochar is sprinkled onto soil it is more likely to suppress soil-derived CO2 emission than when mixed in soil; however, the trend based on biochar quality was unclear.

© 2019 by the authors. Few studies have reported the estimation of nitrogen (N) deposition, including dissolved organic N (DON) fluxes, through water flows and the contribution of snowfall in Asia. In this study, the concentrations and fluxes of DON and dissolved inorganic N (DIN) in bulk precipitation (BP), the throughfall (TF) of trees and understory dwarf bamboo, and stemflow (SF) were evaluated in a cool-temperate forest over three years to clarify N fluxes via precipitation and responses of trees and understory canopies to N deposition. The input of N to the study site in BP was 11.1 ± 1.71 kg N ha-1 year-1, with a significant contribution from DON (78%). Snowfall fluxes contributed up to 46% of the totalNinput, with variations related to the amount of snowfall (2.08-5.52 kgNha-1 year-1). The forest canopy enriched DON (2.11 ± 0.42 kg N ha-1 year-1) but consumed NO3 + NO2-N (-0.73 ± 0.19 kg N ha-1 year-1). In contrast, through the understory bamboo canopy, DON (-1.02 ± 0.55 kg N ha-1 year-1) decreased while DIN (0.35 ± 0.44 kg N ha-1 year-1) increased. This study indicates that DON and snowfall should not be neglected when evaluating total N deposition into forest ecosystems, especially in remote regions. The canopy processes related to the dissolved N in the presence of understory plants might have significant implications for the internal N cycle in forest ecosystems.

© 2019 Elsevier Ltd Dissolved organic matter (DOM) plays an important role in sustaining ecosystem services of mangrove forests through well-described biogeochemical and ecological functions. This study was conducted in the Fukido River (Ishigaki Island, Japan) to better understand the seasonal and episodic changes in DOM concentration and composition in a subtropical mangrove system. Water samples were collected seasonally along a headwater–mangrove–sea transect on 10 occasions from September 2014 through June 2016. DOM was fractionated based on hydrophobicity into two fractions (hydrophobic and hydrophilic) and also analyzed by excitation-emission matrix spectroscopy combined with parallel factor analysis (PARAFAC). Although seasonal changes in DOM concentration and composition were not observed, both hydrophobic and hydrophilic DOM concentrations and levels of the identified three PARAFAC components clearly increased during a typhoon event. It is suggested that episodic increases in freshwater input due to a typhoon caused enhanced leaching of DOM from mangrove litter and dissolution of mangrove soil organic matter (SOM), which was otherwise retained in the mangrove soil by salinity-induced aggregation. The aggregation–dissolution properties of SOM are crucial in determining the magnitude of DOM outwelling and possibly SOM accumulation rate by enhancing advective DOM exchanges. Future studies are needed to evaluate the size of the carbon pool and outwelling of DOM after classifying mangrove forests based on the hydrological regime that influences biogeochemical conditions in the forests. Regional Index Terms: Japan, Okinawa, Ishigaki, Fukido.

© 2019 Elsevier B.V. There are insufficient data regarding forest productivity of mature mangroves near the northern limit of mangrove distribution in subtropical East Asia. We conducted the present study to determine the stand dynamics and net primary production (NPP) of a mature mangrove forest on Ishigaki Island in south-western Japan over three successive years, using a large permanent plot. The two mangrove species present were Bruguiera gymnorrhiza and Rhizophora stylosa. The aboveground biomass in the plot was high, despite the high latitude (24 ∘ 29 ′ N) of the site, gradually increasing from 158.4 Mg ha −1 to 164.6 Mg ha −1 over the three-year study period. This increase was attributable to an increase in the biomass of B. gymnorrhiza. The biomass of R. stylosa, however, decreased during the same period because of a threefold-higher mortality rate and a lower relative growth rate of this species compared with B. gymnorrhiza. The mangrove forest was in the late successional stage, following a pioneer R. stylosa forest that was widely distributed throughout the study plot as standing dead trees and logs. The aboveground NPP of the mangrove forest was 10.66 ± 1.46 Mg ha −1 y −1 , partitioned into 3.10 ± 0.51 Mg ha −1 y −1 as woody NPP (net increase in aboveground woody parts, SI), and 7.56 ± 0.99 Mg ha −1 y −1 as foliage NPP (litter production, including foliage and reproductive organs, L n ). The mature mangrove forest had a relatively low SI/L n ratio (0.41 ± 0.03), although litter fall production was within the range previously recorded for mangroves. The lower woody NPP in the mature mangroves was due to the exclusion of R. stylosa once the stand was in a late successional stage dominated by B. gymnorrhiza.

© 2019, © 2019 Japanese Society of Soil Science and Plant Nutrition. The role of biochar in the mitigation of CO2 emissions has been extensively studied in agricultural soils but is not well understood in Japanese forest soils, especially in relation to CO2 emissions from applied biochar and native soil C (i.e., the priming effect; PE). We hypothesized that the type of biochar and/or the application method (mixed or sprinkled) affect the direction and magnitude of PE in forest soil, and in particular, negative PE can be achieved relatively easily if biochar produced under higher temperature conditions were sprinkled on the soil surface. To test our hypothesis, we measured CO2 emissions from biochar-amended brown forest soil in Japan and examined its PE by conducting a medium-term (~4 months) incubation study. As substrates, we used plain straw from the C4 grass Miscanthus sinensis (SU) and two qualities of biochar produced from it at either 300°C (BC300) or 800°C (BC800) and compared two application methods: mixed into or sprinkled onto the soil. BC800 had a greater C content and C:N ratio as well a lower volatile matter content and higher nonvolatile matter content than BC300

Humic substances (HS) are the primary constituents of dissolved organic matter (DOM) and play pivotal roles in aquatic systems. Optical indices of DOM, such as specific UV absorbance (SUVA254), the fluorescence index (FI) and biological index (BIX), have gained wide interest because of their ease of use. In this study, we explored the relationship between HS and the indices in the Trat River Basin (eastern Thailand) from headwaters to the river mouth through the distinct dry and rainy seasons to examine whether changes in index values reflect variability in the relative contribution of HS to DOM, or %HS. The results show that %HS and the indices did not exhibit significant linear relationships (FI and BIX, P &gt
0.05), or the relationships changed seasonally (SUVA254). However, analyzing the indices versus %HS did show clear DOM composition changes by season with more humic-like or terrestrial material in the rainy season. Relationships between DOM and dissolved iron (dFe) concentrations were also explored. Separating the relationships of DOM versus dFe into HS versus dFe and non-HS versus dFe provides us the opportunity to better understand which fraction contributes more to dFe mobilization. The results indicate stronger positive linear relationships between HS and dFe concentrations independent of river tributary. Overall, this study highlights the importance of quantifying HS for the study of DOM dynamics or compositional changes along a river transect as well as for DOM-induced iron mobilization.

© 2018 Cambridge University Press. The variation in CO2 flux from the forest floor is important in understanding the role of mangrove forests as a carbon sink. To clarify the effects of soil temperature and tidal conditions on variation in CO2 flux, sediment-atmosphere CO2 fluxes were measured between June 2012 and May 2013. We used the closed chamber method for two plots, with a 0.5 m difference in elevation (B, high elevation; R-B, low elevation), in a mangrove forest in south-western Japan. CO2 fluxes were highest in the warm season and showed a weak positive correlation with soil temperature in both forests. Estimated monthly CO2 flux showed moderate seasonal variation in accordance with the exposure duration of the soil surface under tidal fluctuation. Additionally, measured CO2 flux and soil temperature were slightly higher in the R-B plot than the B plot, although estimated annual CO2 flux was higher in the B plot than the R-B plot due to different exposure durations. These results suggest that variation in the exposure duration of the forest floor, which changes seasonally and microgeographically, is important in evaluating the annual CO2 flux at a local scale and understanding the role of mangrove ecosystems as regulators of atmospheric CO2.

We report long-term continuous phenological and sky images taken by time-lapse cameras through the Phenological Eyes Network (http://www.pheno-eye.org. Accessed 29 May 2018) in various ecosystems from the Arctic to the tropics. Phenological images are useful in recording the year-to-year variability in the timing of flowering, leaf-flush, leaf-coloring, and leaf-fall and detecting the characteristics of phenological patterns and timing sensitivity among species and ecosystems. They can also help interpret variations in carbon, water, and heat cycling in terrestrial ecosystems, and be used to obtain ground-truth data for the validation of satellite-observed products. Sky images are useful in continuously recording atmospheric conditions and obtaining ground-truth data for the validation of cloud contamination and atmospheric noise present in satellite remote-sensing data. We have taken sky, forest canopy, forest floor, and shoot images of a range of tree species and landscapes, using time-lapse cameras installed on forest floors, towers, and rooftops. In total, 84 time-lapse cameras at 29 sites have taken 8 million images since 1999. Our images provide (1) long-term, continuous detailed records of plant phenology that are more quantitative than in situ visual phenological observations of index trees
(2) basic information to explain the responsiveness, vulnerability, and resilience of ecosystem canopies and their functions and services to changes in climate
and (3) ground-truthing for the validation of satellite remote-sensing observations.

Rapid glacial retreat in the High Arctic causes the expansion of new habitats, but the successional trajectories of soil microbial communities are not fully understood. We examined microbial succession along a chronosequence twice with a 10-year interval in a High Arctic glacier foreland. Soil samples were collected from five study sites with different ages and phospholipid fatty acids analysis was conducted to investigate the microbial biomass and community structure. Microbial biomass did not differ significantly between the two sampling times but tended to increase with the chronosequence and showed a significant correlation with soil carbon (C) and nitrogen (N) content. Microbial community structure clearly differed along the chronosequence and was correlated with C and N content. The largest shift in community structure over 10 years was observed in the newly exposed sites after deglaciation. The accumulation of soil organic matter was regarded as an important determinant both of microbial biomass and community structure over the successional period. In contrast, the initial microbial community on the newly exposed soil changed rapidly even in the High Arctic, suggesting that some key soil processes such as C and N cycling can also shift within the relatively short period after rapid glacial retreat.

The chamber method with plant clipping has been widely used for measuring soil respiration (SR) in grassland ecosystems. However, plant clipping may cause overestimation of SR by changing the environmental factors and injuring the plants. To solve these problems, we developed a new non-destructive method using multiple-microchambers (3 cm diameter, 8 cm height), which enables measurement of SR without plant clipping by installing chambers into gaps among the grasses. The new method was compared with the conventional method at various flow rates in vitro to assess the accuracy of SR measurement. The new method overestimated the SR rate
however, the ratio of overestimation to the conventional method was constant for each flow rate. These ratios fitted the logarithmic curve, indicating the potential for correction of the SR rate measured by the new method using the logarithmic equation. The corrected SR rate obtained by the new method was equal to the rate by the conventional method. This suggests that accurate measurement of SR in grassland ecosystems is possible using the multiple-microchambers method. We then compared the non-destructive method and the destructive method in situ on summer season and found that the destructive method overestimated SR rate in the grassland ecosystem by about 276% on average. There were two possible reasons for this overestimation
first, the clipping treatment may change environmental conditions such as soil temperature and soil water content, and second, it may directly increase plant respiration.

Secondary lucidophyllous forest is one of the dominant forests in human-dominated subtropical/warm-temperate regions in East Asia. There were few direct monitoring techniques to elucidate the following hypotheses: (a) self-thinning may govern the stand development process and (b) wood production decline can be observed during secondary succession in a lucidophyllous forest. We conducted a long-term study at a permanent plot in central Japan, since 1989. The forest consists mainly of Castanopsis cuspidata in a canopy layer, Cleyera japonica, and Eurya japonica in a subtree layer. During the 28-year period, the basal area of the stand significantly increased due to the growth of C. cuspidata, from 29.18 +/- 1.84 (87.8% of total) to 38.71 +/- 2.22 m(2) ha(-1) (91.9%), while the stem density of C. cuspidata significantly decreased from 666 +/- 13 to 404 +/- 10 stems ha(-1) in proportion to accumulating biomass (117.8 to 166.6 ton ha(-1)). The annual woody net primary production ranged from 2.40 +/- 0.13 to 3.93 +/- 0.33 ton ha(-1) year(-1) as a nearly 70-year-old forest. There was no age-related decline of woody net primary production (NPP) was found during secondary succession, and the growth of individual tree still increased when the self-thinning process governed the stand.

Dissolved organic carbon (DOC) plays an important role in C cycling in forest ecosystems. Here we measured the concentrations and fluxes of DOC in a cool-temperate broad-leaved deciduous forest (Takayama Forest) to quantify the contribution of DOC from different forest water flux conditions. Mean DOC concentration during the growing season increased in the sequence from bulk precipitation (2.98 +/- 0.45 mg L-1), throughfall above dwarf bamboo (6.84 +/- 0.45 mg L-1), throughfall below dwarf bamboo (7.08 +/- 0.42 mg L-1), stemflow (15.05 +/- 0.98 mg L-1), and litter leachate (21.33 +/- 1.01 mg L-1). Litter leachate DOC concentration, being high in spring and autumn, which was fairly correlated with the amount of litterfall of bamboo and trees. In stemflow, the DOC concentration was high during early summer and gradually decreased, in addition, it also showed dramatic variation among different plant species. Litter leachate (72.5%) accounted for most of the DOC input to the soil during the growing season (311.5 kg C ha(-1) 7 months(-1)), while stemflow (1.6%) contributed the least. A great quantity of precipitation at the study site was associated with a subsequent high atmospheric contribution of DOC flux (8.6%), which was more than half of throughfall (16.5%). The high input of DOC to the soil and andisol soil characteristics at the Takayama Forest suggest that the DOC fluxes are vital to the soil carbon sequestration. Therefore, DOC fluxes should be taken into account when the carbon balance is assessed at forest ecosystems.

This paper reports seasonal data regarding leaf litter for 14 deciduous broad-leaved species and one evergreen coniferous species as well as leaf area index (LAI) data for the 14 deciduous broad-leaved species in a cool-temperate deciduous broad-leaved forest in Japan. The seasonal leaf biomass of various tree species is important for accurately evaluating ecosystem functions such as photosynthesis and evapotranspiration under climate change. However, there is a lack of freely available, long-term data. We collected litterfall every 1 to 4 weeks from September or October to November or December each year from 2005 to 2014 in Takayama, Japan (36A degrees 08'46aEuro(3)N, 137A degrees 25'23aEuro(3)E, 1420 m a.s.l.). After sorting the litter into leaves (according to species categories), stems + branches, and "other", we dried and weighed the litter groups. We also collected seasonal leaf data (number of leaves and leaf length and width) for each broad-leaved species, which we recorded every 1 to 4 weeks from April or May to October or November using multiple target shoots. To estimate the LAI in autumn for each deciduous broad-leaved species, we used a semi-empirical model of the vertical integration of leaf dry mass per unit leaf area. To estimate the LAI in spring and summer, we used the relationship between the LAI in autumn and the seasonal leaf data. Our data provide input, calibration, and validation parameters for determining LAI based on satellite remote-sensing observations or radiative transfer models and for use in ecosystem models.

The structural complexity, especially canopy and gap structure, of old-growth forests affects the spatial variation of soil respiration (Rs). Without considering this variation, the upscaling of Rs from field measurements to the forest site will be biased. The present study examined responses of Rs to soil temperature (Ts) and water content (W) in canopy and gap areas, developed the best fit model of Rs and used the unique spatial patterns of Rs and crown closure to upscale chamber measurements to the site scale in an old-growth beech-oak forest. Rs increased with an increase in Ts in both gap and canopy areas, but the effect of W on Rs was different between the two areas. The generalized linear model (GLM) analysis identified that an empirical model of Rs with the coupling of Ts and W was better than an exponential model of Rs with only Ts. Moreover, because of different responses of Rs to W between canopy and gap areas, it was necessary to estimate Rs in these areas separately. Consequently, combining the spatial patterns of Rs and the crown closure could allow upscaling of Rs from chamber-based measurements to the whole site in the present study.

We measured the CO2 flux from sediment surfaces (soil respiration in the pedosphere to the atmosphere) in a mangrove forest using an improved automatic open/close chamber (AOCC) method. Soil respiration rates and environmental factors were continuously measured from 4 to 8 July, 2013, in a mangrove dominated by Bruguiera gymnorrhiza. Variation in respiration rate did not exhibit a clear correlation with soil temperature. However, tidal effects were related to variation in soil temperature and may also have contributed to variation in respiration rate. High respiration rates were detected immediately before submergence or after exposure, due to the physical effects of tidal variation. Respiration rates during the period of exposure were lower than those in terrestrial ecosystems, likely due to three factors unique to mangrove forests: soil respiration measurements generally do not include root respiration, organic matter decomposition is restricted to a shallow anaerobic area, and some mineralized carbon is lost as dissolved inorganic carbon. Respiration rates during submergence were half of those measured during exposed conditions, suggesting that previous studies overestimated annual soil respiration. Therefore, measuring soil respiration rates during both exposed and submerged conditions using the AOCC method provides a much more accurate understanding of carbon dynamics in the pedosphere of mangrove forests.

To clarify the effects of long-term warming on ecosystem matter cycling, we conducted an in situ 7-year experimental warming (2009-2015) using infrared heaters in a cool temperate semi-natural grassland in Japan. We measured plant aboveground biomass, soil total C and N, soil inorganic N (NH4 (+)-N and NO3 (-)-N), and soil microbial biomass for 7 years (2009-2015). We also measured heterotrophic respiration for 2 years (2013-2014) and assessed net N mineralization and nitrification in 2015. We found that warming immediately increased plant aboveground biomass, but this effect ceased in 2013. However, the soil microbial biomass was continuously depressed by warming. Soil inorganic N concentrations in warmed plots substantially increased in the later years of the experiment (2013-2015) and the potential net N mineralization rate was also higher than in the earlier years. In contrast, heterotrophic respiration decreased with warming in 2013-2014. Our observations indicate that long-term warming has a contrasting effect on plants and soil microbes. In addition, the warming could have different effects on subterranean C and N cycling. To enhance the accuracy of estimation of future climate change, it is essential to continuously observe the warming effects on ecosystems and to focus on the change in subterranean C and N cycling.

We examined two hypotheses based on laboratory amendment experiments: (1) that development of the soil microbial community on volcanic deserts was regulated by substrate limitation; and (2) that the type and the extent of substrate limitation would change along the succession gradient. Soils were collected from the early (Stage B) and the late (Stage F) stages of primary succession of a subalpine volcanic desert on Mt. Fuji and they were amended with three carbon (C) sources (glucose, cellulose, or lignin), inorganic nitrogen (N), or phosphorus (P) sources alone or in a mixture. Respiration rates were monitored for 25 days and changes in microbial biomass and community structure were examined using the content and composition of phospholipid fatty acids. For both soils, the magnitude of the microbial response differed depending on the type of C source and it decreased in the following order reflecting the availability to microorganisms: glucose > cellulose > lignin. For Stage B soil, although any single amendment did not affect the microbial properties, combined amendment of C (glucose) and N increased microbial respiration and biomass and shifted the microbial community structure. In contrast, microbial properties in Stage F soils responded positively to single amendments of C source. Our results suggest that the microbial community in the early stage of succession is primarily limited by simultaneous shortage of C and N sources but the quality of the C source becomes more important in the late successional stages, which have large, but recalcitrant, organic matter pools in the soil. (C) 2016 Elsevier Masson SAS. All rights reserved.

We evaluated the uncertainty in the estimation of year-to-year variability in the timing of leaf fall detected by the analysis of red, green and blue (RGB) values extracted from daily phenological images in a deciduous broad-leaved forest in Japan. We examined (1) the spatial distribution of individual tree species within a 1-ha permanent plot and the spatio-temporal variability of leaf litter of various species for 8 years; and (2) the relationship between the year-to-year variability of leaf fall detected by leaf litter and that detected by phenological images of various species. Uncertainties were caused by (1) the heterogeneous distribution of each species within the whole forest community; (2) the year-to-year variability of the timing of leaf fall among species; and (3) differences in leaf colouring and leaf fall patterns among species. Our results indicate the importance of integrating RGB analysis of each species and of the whole canopy on the basis of spatial locations of individuals and proportions of tree species within a forest to reduce uncertainty. (C) 2015 Elsevier B.V. All rights reserved.

Soil respiration (R (s) ) is a key component in the estimation of the net ecosystem production (NEP) of old-growth forests, which are generally thought to have ceased carbon accumulation. The objectives of the present study were to characterize the spatial and temporal patterns of R (s) , and to identify the determinants of the spatial and temporal variability of R (s,) using general linear mixed models (GLMM), in an old-growth beech-oak forest. GLMM analyses identified monthly effect as a significant explanatory variable for temporal variation, as well as gap/canopy and soil water content (SWC) as explanatory variables for spatial variation, in R (s) . The complexity of vertical structure in the forest was reflected in the spatial pattern of R (s) , which was higher in canopy areas than in gap areas during the growing season, except in November. This spatial pattern was not affected by soil temperature. Moreover, SWC did not differ between gap and canopy areas, although SWC partially explained the spatial heterogeneity in R (s) . The carbon:nitrogen ratios of soil organic matter in canopy areas were significantly higher than those in gap areas. Fine root biomass was 1.7-fold greater in canopy areas than in gap areas, likely because of the higher R (s) in canopy areas, and root respiration made a much large contribution to R (s) than heterotrophic respiration. The different patterns of fine root biomass between gap and canopy areas mainly control the spatial heterogeneity in R (s) ; thus, it is worth considering the gap/canopy variability in R (s) when evaluating annual efflux in old-growth forests.

Moss tundra that accumulates a thick peat layer is one of the most important ecosystems in the High Arctic, Svalbard. The importance of this ecosystem for carbon sequestration was estimated from the apparent rates of carbon accumulation based on the C-14 age and amount of peat in the active layer. The study site at Stuphallet, Brogger Peninsula, northwestern Svalbard was covered with a thick peat layer dominated by moss species such as Calliergon richardsonii, Paludella squarrosa, Tomenthypnum nitens, and Warnstorfia exannulata. The average thickness of the active layer (brown moss and peat) was approximately 28 cm in 1 August 2011. The calibrated (cal) age of peat from the bottom of the active layer (20-30 cm below the peatland surface) ranged from 81 to 701 cal yr BP (median value of 2s range). Based on the total carbon (4.5-9.2 kg C m(-2)), the apparent rate of carbon accumulation in the active layer was 9.0-19.2 (g C m(-2) yr(-1)), which is similar to or greater than the net ecosystem production or net primary production reported for other vegetation types in this area. Our data suggest that moss tundra plays an important role in carbon sequestration in this area. (C) 2015 Elsevier B.V. and NIPR. All rights reserved.

To assess soil microbial response to global warming in cool temperate semi-natural grassland, we conducted an in situ warming experiment in grassland located in the mountains of central Japan. Five pairs of plots (control and warmed) of Zoysia japonica were established. For each pair of plots, one was warmed by ca. 2 A degrees C using infrared heaters during the growing seasons of 2009-2011. Above-ground biomass of Z. japonica was estimated using the modified point-frame method. Soil organic matter contents, soil total carbon and nitrogen contents, as well as inorganic nitrogen (ammonium and nitrate) contents were determined from soil samples. Total phospholipid fatty acid (PLFA) contents and PLFA compositions were determined and used as indices for total microbial biomass and community structure, respectively. Analyses showed that the warming increased the above-ground biomass of Z. japonica significantly. Soil organic matter and soil total nitrogen contents were significantly decreased, while soil ammonium content was significantly increased in the warmed plots. Soil microbial biomass (especially fungal biomass) was lower in the warmed plots, probably reflecting higher temperature, lower soil water content, and/or depletion in available nutrients. The significant decrease in fungal biomass, in combination with our PLFA composition data, suggests that the soil microbial community structure shifted from a fungal-dominated to a bacteria-dominated one, causing changes in community-level physiological activity.

Soil organic layer samples of two different forest types were observed using compact MRI to visualize internal structure and clarify physical properties of forest soil. Soil pores and organic materials were distinguished by differences in proton mobility and visualized with a spatial resolution of 234 A mu m. Soil pore ratios and water mobility were calculated by image processing, and their differences between the two forest soils were detected. Our results suggest that compact MRI has potential for non-destructive analysis of soil physical properties and is expected to have significant applications in ecological studies.

Long-term continuous phenological observation of cherry tree blooming is an important and challenging task in the evaluation of year-to-year weather and climate changes in spring in Japan. Here, (1) we performed daily field observations with a time-lapse digital camera in a deciduous broad-leaved forest in Japan from January 2004 to December 2013; and (2) we detected year-to-year variations in the blooming phenology of Prunus sargentii by visual inspection of the images and by image analysis. We found that (1) the red digital numbers (i.e., the digital intensity values of the red pixels; DNR) extracted from the time-lapse digital camera images tended to peak in full bloom; and (2) the green excess index (GET) (based on the red, green, and blue digital numbers) tended to be lowest in full bloom. These results indicate that DNR and GEI are useful for detecting the timing of full bloom in P. sargentii.

Livestock dung provides an important direct pathway by which carbon and nutrients enter soils in pasture ecosystems and affects carbon and nitrogen cycling indirectly through changes in soil and plant properties. Here, we quantify dung deposition, decomposition, and the effects of dung on soil and plants in a Zoysia japonica grassland in Japan. We determined (1) the distribution of dung, (2) the mass loss rate of dung and the amount of carbon respired as CO2, and (3) changes in soil properties and aboveground biomass of Z. japonica. Dung deposition was 4.0-9.7 g C and 0.4-1.0 g N m(-2) year(-1) and distributed patchily (Morishita's I (delta) > 1). Most (71 %) of the carbon in dung deposited in June was lost within a single grazing period by aerobic decomposition, more than mass loss rate of Z. japonica litter in the first year (about 50 %), suggesting that grazing and defecation can accelerate carbon cycling compared with the typical litterfall-decomposition regime. Nitrogen in dung mass entered the soil as ammonium nitrogen and was nitrified. The spatiotemporal distribution of these processes corresponded to that of stimulated Z. japonica growth. These results suggested that dung deposition significantly affected the inorganic nitrogen status of soil and, therefore, the growth of Z. japonica. However, these effects were very restricted temporally (July-August) and spatially (within 10 cm from dung edge). Thus, such spatiotemporally restricted effects combined with the patchy distribution of dung may contribute to the heterogeneous structure of pasture ecosystems.

Mud shrimps, Upogebia spp., are major constituents of macrobenthic communities in tidal flats in Japan. The impact of Upogebia yokoyai on carbon flow on tidal flats was examined by comparing CO2 emission rates from plots with and without burrows in the Kurose River estuary, Japan. In situ CO2 emission rates from plots with burrows were significantly higher than from those without. Laboratory measurements using sediment core samples that excluded respiration of macrobenthic organisms showed similar trends. Although there were no significant differences in grain size distribution, water content, or ignition loss between the sediment cores with and without burrows, oxidation-reduction potential was significantly higher in sediment cores with burrows. Analysis of phospholipid fatty acids (PLFA) indicated that microbial biomass and community structure did not differ significantly between cores with and without burrows. However, microbial respiration activity, as indicated by CO2 emission rates per total PLFA content, was significantly higher in sediment cores with burrows than in those without. Our results indicate that burrows of U. yokoyai change the physicochemical conditions and increase microbial activity in the sediment, significantly affecting carbon flow in the tidal flat.

Trenching (Tr), root biomass regression (RR), and root excising (RE) methods were used to estimate the contribution of root (RR) and heterotrophic (HR) respiration to soil respiration (SR) in a cool-temperate deciduous forest in central Japan. The contribution ratios of RR to SR were 23 % (-16 to 46 %), 11 % (-19 to 61 %), and 115 % (20 to 393 %), as estimated by the Tr, RR, and RE methods, respectively. The contribution ratio showed clear seasonal variation with high values in summer for the Tr method, while they were undetectable for the RR and RE methods because of some methodological problems. These results suggest the Tr method is the best of the three methods used to estimate the contribution ratio of RR and HR to SR in the forest. Annual SR, RR, and HR rates, estimated by the Tr method, were 479, 369, 110 gC m(-2) year(-1), respectively. The seasonal variation of SR was mainly influenced by HR (77 %) throughout the year, while the influence of RR on SR was strongest in summer (46 %). This effect occurred because RR (Q (10) = 7.5) is more sensitive to temperature than HR (Q (10) = 3.2). Also, the contribution of fine RR to total RR was higher than that of coarse RR because of high respiratory activity (Q (10) and R (10)) as well as the large biomass of fine roots. These results suggest that each component of SR responds differently to the same environmental factors and their relative influence on SR changes across the seasons.

To study the relationship between vegetation development and changes in the soil microbial community during primary succession in a volcanic desert, we examined successional changes in microbial respiration, biomass, and community structure in a volcanic desert on Mount Fuji, Japan.
Soil samples were collected from six successional stages, including isolated island-like plant communities. We measured microbial respiration and performed phospholipid fatty acid (PLFA) analysis, denaturing gradient gel electrophoresis (DGGE) analysis, and community-level physiological profile (CLPP) analysis using Biolog microplates.
Microbial biomass (total PLFA content) increased during plant succession and was positively correlated with soil properties including soil water and soil organic matter (SOM) contents. The microbial respiration rate per unit biomass decreased during succession. Nonmetric multidimensional scaling based on the PLFA, DGGE, and CLPP analyses showed a substantial shift in microbial community structure as a result of initial colonization by the pioneer herb Polygonum cuspidatum and subsequent colonization by Larix kaempferi into central areas of island-like communities. These shifts in microbial community structure probably reflect differences in SOM quality.
Microbial succession in the volcanic desert of Mt. Fuji was initially strongly affected by colonization of the pioneer herbaceous plant (P. cuspidatum) associated with substantial changes in the soil environment. Subsequent changes in vegetation, including the invasion of shrubs such as L. kaempferi, also affected the microbial community structure.

The distribution of organic carbon and its relationship to vegetation development were examined on a glacier foreland near Ny-angstrom lesund, Svalbard (79 degrees N). In a 0.72-km(2) area, we established 43 study plots on three line transects along primary succession from recently deglaciated area to old well-vegetated area. At each plot, we measured the type and percent coverage of vegetation types. The organic carbon content of vegetation, organic soil, and mineral soil samples was determined based on their organic carbon concentration and bulk density. Cluster analysis based on vegetation coverage revealed five types of ground surfaces representing variations in the amounts and allocation patterns of organic carbon. In the later stages of succession, 7%-24% and 31%-40% of organic carbon was contained in the organic and deeper soil layers, respectively. Organic carbon storage in the later stages of succession ranged from 1.1-7.9 kg C m(-2). A larger amount of organic carbon, including ancient carbon in a raised beach deposit, was expected to be contained in much deeper soil layers. These results suggest that both vegetation development and geological history affect ecosystem carbon storage and that a non-negligible amount of organic carbon is distributed in this High Arctic glacier foreland. (C) 2011 Elsevier B.V. and NIPR. All rights reserved.

Polar willow (Salix polaris Wahlenb.), a mycorrhizal dwarf shrub, colonizes recently deglaciated areas in the High Arctic, Svalbard. To clarify successional changes in ECM fungi associated with S. polaris after glacier retreat, we examined the diversity and density of ECM fungi in culture and field conditions. Plant and soil samples were collected from three sites of different successional stages in the deglaciated area of Austre Broggerbreen, near Ny-lesund, Svalbard. The successional stages were early stage with newly exposed bare ground (site I), transient stage with scattered colonization of Salix (sites IIa and IIb), and late stage with well-developed vegetation (site III). No ECM colonization on Salix was observed in soils collected from bare ground in early and transient stages (sites I and IIa). However, most Salix individuals showed ECM colonization in soils collected from sites close to Salix colonies in transient and late stages (sites IIb and III). Based on molecular analyses and operational taxonomic unit (OTU: > 95% ITS sequence similarity) delimitations, we identified 15 OTUs/species in eight genera. The dominant OTU/species of ECM fungi identified in the transient and late stages was Geopora sp.1 and Cenococcum sp.1, respectively. In the culture experiment, ECM diversity was greater in late stage (eight OTUs/species) than in transient stage (three OTUs/species). This pattern was consistent with field observations, i.e., late-stage sites contained more OTUs/species of ECM fungi. These results indicate that species diversity of ECM fungi increases and the dominant species changes with the progress of succession after glacier retreat in the High Arctic.

The polar willow (Salix polaris), predominant in the late successional stage in deglaciated areas of Ny-Alesund, Svalbard, is rarely found in the early stage, when purple saxifrage (Saxifraga oppositifolia) dominates. To elucidate the pattern and the mechanism of successional change from the Saxifraga stage to the Salix stage, we examined the distribution pattern, size structure and habitat conditions of a colonizing Salix population in the seral stage where Salix was invading Saxifraga-dominated sites. The present distribution pattern and aerial photographs taken in the past suggest that Salix colonization at this site commenced within the last 70 years. We found 115 Salix individuals (22 male, 13 female and 80 unknown) in a 30 m 30 m quadrat on the seral stage. Although the largest individual had a size of 2000 cm(2) (length x maximum width), the majority (84%) of individuals were smaller than 100 cm(2). The seedling size distribution, as inferred from the leaf scar number, indicated that annual recruitment was slight. Of the individuals observed about 75% had colonized bare ground; only four individuals grew within Saxifraga colonies. No significant difference was found in soil characteristics (water content, and carbon and nitrogen concentrations) between the seral stage and the earlier stage prior to colonization by Salix. These results suggest that difficulties in seed production, germination and/or seedling establishment of Salix, rather than soil formation by preceding species (Saxifraga), limits the early-stage colonization by Salix.

We examined the photosynthetic characteristics and net primary production of biological soil crusts to evaluate their contribution to the carbon cycle in the High Arctic glacier foreland.
Biological soil crust samples were collected from a deglaciated area in Ny-Alesund, Svalbard, Norway. Net photosynthetic rates (Pn) and dark respiration rates (R) of biological soil crusts were determined using CO(2) gas exchange rates. We examined the effects of moisture conditions, temperature and photon flux density on Pn and R, and estimated the net primary production by a model based on the relationships between abiotic factors and Pn and R.
The maximum Pn value occurred at 50% of the maximum water-holding capacity. Pn decreased with increasing temperature and dropped below zero at high temperatures (c. > 13 degrees C). The estimated net primary production of the biological soil crust was greater than the net primary production of other vegetation when based on ground surface area, during the early stage of primary succession. Model simulation showed that the net primary production of the biological soil crust decreased with increasing temperature.
These results suggest that biological soil crust productivity plays an important role in the carbon cycle during the early stage of succession of the High Arctic glacier foreland, and is susceptible to temperature increases from global warming.

A significant temperature increase in Arctic tundra ecosystems was observed during 20th century and the temperature increase is expected to be most pronounced in the Arctic at the end of this century. This paper introduces photosynthetic characteristics of three cryptogamic plants (moss, lichen and biological soil crust) on the glacier foreland in the high Arctic, Ny-Alesund, Svalbard, Norway. We constructed a model for estimating the net primary production of the three plants to evaluate an effect of temperature increase on the net primary production.

Raised beach deposits are widespread on the north-western coast of Spitsbergen, Svalbard. To elucidate the importance of these deposits in an ecosystem carbon cycle, we measured the concentrations of organic carbon and adenosine 5-triphosphate (ATP; an index of living microbial biomass) in a raised beach deposit found under terrestrial vegetation in Ny-Alesund. A shell in the deposit found at a depth of ca. 20 cm below the ground surface had a (not calibrated) C-14 age of 11080 +/- 140 yr BP, whereas soil organic carbon in the same deposit showed an older C-14 age (22380 +/- 90 yr BP). Organic carbon concentration in the layer of 20-40 cm, belowground was about 1-2%, which was comparable to those in shallower mineral soil layers. Results of ATP analyses suggested that low but non-negligible amounts of microorganisms existed in the deposit. The proportion of biomass carbon to soil organic carbon tended to decrease with increasing depth, suggesting that organic carbon in the deep layer was less available to microorganisms than that in the shallow layers.

We used phospholipid fatty acid (PLFA) analysis to examine the relation of microbial biomass and community composition to vegetation zonation on a coastal sand dune. Soil samples were collected along 3 line transects established from the shoreline to the inland bush. Total PLFA content and PLFA composition of soils were used as indices of total microbial biomass and community composition, respectively. The microbial biomass was much higher in the inland Vitex rotundifolia zone than in the seaside plots. The microbial community composition also differed among the vegetation zones, with a higher contribution of fungal biomarkers in the inland plots. The microbial biomass increased significantly with increasing soil organic matter (SOM) content, but was not correlated with soil salinity. These results suggest that microbial biomass in the coastal sand dune was controlled primarily by the accumulation of SOM. The microbial community composition also changed with SOM content in the seaside plots, but SOM had little effect in the inland plots. These results suggest that the factors limiting the microbial community composition differed with location on the dune.

The solfatara field is a unique ecosystem characterized by harsh conditions such as acidic soils. We examined the respiration rate and phospholipid fatty acid (PLFA) content of solfatara soils and their responses to carbon and nitrogen addition. to determine whether soil microbial respiration and biomass in a solfatara field are limited by substrate availability. Soil samples were collected from locations along a transect across a solfatara field in Oita Prefecture, Japan. The soil in the central part of the solfatara field was highly acidic (pH 2.4) and contained low amounts of carbon and nitrogen. Low basal respiration rates were detected in these soil samples. Measurements of substrate-induced respiration (SIR) and PLFA contents suggested that it was partly attributable to low microbial biomass. Addition of a carbon source (glucose) to the solfatara soil engendered a marked increase in the microbial respiration rate, whereas the nitrogen source (ammonium nitrate) application had no marked effect. Addition of both carbon and nitrogen caused a nearly eightfold increase in the microbial respiration rate and a threefold increase in the total PLFA contents. These results suggest that some acidophilic and/or acid-tolerant microorganisms exist in solfatara soil, but that their respiration and biomass are limited by low substrate availability. (c) 2006 Elsevier Masson SAS. All rights reserved.

The hypotheses that carbon and nitrogen availability limit microbial activity, and that the key factors limiting microbes vary along the successional gradient were tested in a High Arctic glacier foreland. We examined the responses of the respiration rate and the phospholipid fatty acid content to the addition of carbon and/or nitrogen. Soil samples were collected from the early stage and late stage of primary succession in the foreland of a glacier near Ny-Alesund, Svalbard. The addition of both carbon (glucose) and nitrogen (ammonium nitrate) engendered an increase in the microbial respiration rate in the early stage of succession. In contrast, the addition of either carbon or nitrogen did not increase the microbial respiration rate. In the late stage of succession the addition of carbon alone, as well as the addition of both carbon and nitrogen, increased the microbial respiration rate. However, neither the addition of carbon nor the addition of nitrogen affected the total phospholipid fatty acid content (an index of microbial biomass) for any soil within 15 days of incubation at 10 degrees C. An increase in the respiration rate was therefore attributed to changes in the physiological activities of the microbial community, such as enzymatic activity. Our study suggests that microbial respiration was limited by the low availability of both carbon and nitrogen in the early stage of succession. Thereafter, nitrogen limitation is mitigated.

1

1