Temperature and precipitation effects on δ13C depth profiles in SOM under temperate beech forests

Enrichment of 13C in SOM with soil depth is related to interacting processes influenced by temperature and precipitation. Our objectives were to derive climate effects on patterns of vertical δ13C values of soil organic matter (SOM) while minimizing the effect of confounding variables. We investigated vertical changes in δ13C values of SOM in 1-cm depth intervals in silvicultural mature beech (Fagus sylvatica L.) forest ecosystems in northern Rhineland-Palatinate across gradients of MAT (7.9 to 9.7 °C mean annual temperature) and MAP (607 to 1085 mm mean annual precipitation) in winter 2011. Forest stands (n = 10) were chosen based on data sets provided by the Rhineland-Palatinate Forest Administration so that variations in these gradients occurred while other environmental factors like physico-chemical soil properties, tree species, stand age, exposition and precipitation (for the temperature gradient) or temperature (for the precipitation gradient) did not differ among study sites. From litter down to the mineral soil at 10 cm depth, soil organic carbon (SOC) content decreased (47.5 ± SE 0.1% to 2.5 ± 0.1%) while the δ13C values increased (− 29.4 ± 0.1‰ to − 26.1 ± 0.1‰). Litter of sites under higher MAP/lower MAT had lower δ13C values which was in line with literature data on climate driven plant physiological process. To compare the dimension of the vertical 13C enrichment, δ13C values were regressed linearly against log-transformed carbon contents yielding absolute values of these slopes (beta). Beta values ranged between 0.6 and 4.5 (range of r from − 0.7 to − 1.0; p < 0.01). Due to an assumed decay continuum and similar variations of δ13C values in litter and in 10 cm depth, we conclude that effects on isotope composition in the Oi layer continue vertically and therefore, δ13C values in litter do not solely control beta values. Beta values decreased with increasing MAT (r = − 0.83; p < 0.05). Reduced soil moisture and therefore both, reduced microbial activity and reduced downward transport of microbial cycled DOM (=13C enriched) might be responsible for less pronounced δ13C depth profiles in case of high temperatures. Greater C:N ratios (lower degradability) of the litter under higher temperatures likely contributed to these depth trends. Beta values increased with increasing MAP (r = 0.73; p < 0.05). We found decreasing C:N ratios in the mineral soil that possibly indicates higher decomposition under higher precipitation. Exclusion of the organic layers from linear regressions indicated a stronger impact of MAP on the development of δ13C depth profiles. Our results confirm temperature and precipitation effects on δ13C depth profiles and indicate stronger 13C enrichment under lower MAT/higher MAP. Therefore, time series of vertical δ13C depth profiles might provide insights into climate change effects.

Reduction of cement leakage by sequential PMMA application in a vertebroplasty model

PURPOSE Leakage is the most common complication of percutaneous cement augmentation of the spine. The viscosity of the polymethylmethacrylate (PMMA) cement is strongly correlated with the likelihood of cement leakage. We hypothesized that cement leakage can be reduced by sequential cement injection in a vertebroplasty model. METHODS A standardized vertebral body substitute model, consisting of aluminum oxide foams coated by acrylic cement with a preformed leakage path, simulating a ventral vein, was developed. Three injection techniques of 6 ml PMMA were assessed: injection in one single step (all-in-one), injection of 1 ml at the first and 5 ml at the second step with 1 min latency in-between (two-step), and sequential injection of 0.5 ml with 1-min latency between the sequences (sequential). Standard PMMA vertebroplasty cement was used; each injection type was tested on ten vertebral body substitute models with two possible leakage paths per model. Leakage was assessed by radiographs using a zonal graduation: intraspongious = no leakage and extracortical = leakage. RESULTS The leakage rate was significantly lower in the "sequential" technique (2/20 leakages) followed by "two-step" (15/20) and "all-in-one" (20/20) techniques (p < 0.001). The RR for a cement leakage was 10.0 times higher in the "all-in-one" compared to the "sequential" group (95 % confidence intervals 2.7-37.2; p < 0.001). CONCLUSIONS The sequential cement injection is a simple approach to minimize the risk for leakage. Taking advantage of the temperature gradient between body and room temperature, it is possible to increase the cement viscosity inside the vertebra while keeping it low in the syringe. Using sequential injection of small cement volumes, further leakage paths are blocked before further injection of the low-viscosity cement.

A first chironomid-based summer temperature reconstruction (13–5 ka BP) around 49°N in inland Europe compared with local lake development

Temperature reconstructions for the end of the Pleistocene and the first half of the Holocene based on biotic proxies are rare for inland Europe around 49°N. We analysed a 7 m long sequence of lake deposits in the Vihorlat Mts in eastern Slovakia (820 m a.s.l.). Chironomid head capsules were used to reconstruct mean July temperature (TJuly), other proxies (diatoms, green algae, pollen, geochemistry) were used to reconstruct local environmental changes that might have affected the climate reconstruction, such as epilimnetic total phosphorus concentrations (TP), lake level changes and development of surrounding vegetation. During the Younger Dryas (YD), temperature fluctuated between 7 and 11 °C, with distinct, decadal to centennial scale variations, that agree with other palaeoclimate records in Europe such as δ18O content in stalagmites or Greenland ice cores. The results indicate that the site was somewhat colder than expected from the general south-to-north YD temperature gradient within Europe, possibly because of north-facing exposition. The warmer phases of the YD were characterised by low water level or even complete desiccation of the lake (12,200-12,400 cal yr BP). At the Late-Glacial/Holocene transition TJuly steeply increased from from 11 to 15.5 °C (11,700-11,400 cal yr BP) - the highest TJuly for entire sequence. This rapid climate change was reflected by all proxies as a compositional change and increasing species diversity. The open woodlands of Pinus, Betula, Larix and Picea were replaced by broad-leaved temperate forests dominated by Betula, later by Ulmus and finally by Corylus (ca 9700 cal yr BP). At the same time, input of eroded coarse-grained material into the lake decreased and organic matter (LOI) and biogenic silica increased. The Early-Holocene climate was rather stable till 8700 cal yr BP, with temporary decrease in TJuly around 11,200 cal yr BP. The lake was productive with a well-developed littoral, as indicated by both diatoms and chironomids. A distinct decline of TJuly to 10 °C between 8700 and 8000 cal yr BP was associated with decreasing chironomid diversity and increasing climate moistening indicated by pollen. Tychoplanktonic and phosphorus-demanding diatoms increased which might be explained by hydrological and land-cover changes. Later, a gradual warming started after 7000 cal yr BP and representation of macrophytes, periphytic diatoms and littoral chironomids increased. Our results suggest that the Holocene thermal maximum was taking place unusually early in the Holocene at our study site, but its timing might be affected by topography and mesoclimate. We further demonstrated that temperature changes had coincided with variations in local hydrology

A new modern analogue reference dataset and its application to the 430-kyr pollen record from Lake Biwa

This study presents a newly compiled dataset of modern pollen and climate data from 798 sites across Japan and the Russian Far East. This comprehensive reference dataset combined with the modern analogue technique (MAT) provides a powerful tool for pollen-based reconstruction of the Quaternary Northwest Pacific climate. Pollen-derived reconstruction of the modern climate at the reference pollen-sampling sites matches well with the estimated modern climate values (R2 values vary between 0.79 and 0.95, and RMSEP values vary between 5.8 and 9.7% of the modern climatic range for all nine tested variables). The successful testing of the method encourages its application to the fossil pollen records. We used a coarse-resolution pollen record from Lake Biwa to reconstruct glacial-interglacial climate dynamics in central Japan since ~438 kyr and compared it to the earlier reconstruction based on a less representative reference dataset. The current and earlier results consistently demonstrate that the coldest glacial intervals experienced pronounced cooling in winter and moderate cooling in summer, supporting the growth of cool mixed forest (COMX) where warm mixed forest (WAMX) predominates today. During the last glacial, maximum (~24 kyr BP) mean temperatures of the coldest (MTCO) and warmest (MTWA) month were about -13 °C (RMSEP = 2.34 °C) and 21 °C (RMSEP = 1.66 °C) respectively, and annual precipitation (PANN) was about 800 mm (RMSEP = 158.06 mm). During the thermal optimums of the interglacial intervals, the temperatures of the coldest and warmest month were above 0 °C and 25 °C respectively, leading to the reconstruction of WAMX and temperate conifer forest (TECO). Although both these vegetation types grow in the southern part of Japan today, WAMX requires warmer space. The presence of WAMX during marine isotope stages (MIS) 11 and 1, and its absence during MIS 9 and MIS 5 contradict the marine isotope and Antarctic ice records, suggesting that the latter two interglacials were the warmest of the last 800 kyr. The apparent contradiction allows at least three different explanations including low temporal resolution of the pollen record; different trends in CO2 concentrations during 'short' and 'long' interglacials; and regional climate variability and non-linear response of different regions to the global forcing. More definitive conclusions will be possible on the basis of forthcoming high-resolution pollen records from central Japan.

Reference datasets and pollen record from sediment core YAK, Yakumo, SW Hokkaido

This study reconstructs middle and late Holocene vegetation and climate dynamics in the Oshima Peninsula, SW Hokkaido, using the published method of biome reconstruction and modern analogue technique applied to the Yakumo pollen record (42°17'03''N, 140°15'34''E) spanning the last 5500 years. Two previously published matrices assigning Japanese plant/pollen taxa to the major vegetation types (biomes) are tested using a newly compiled dataset of 78 surface pollen spectra from Hokkaido. With both matrices showing strengths and weaknesses in reconstructing cool mixed and temperate deciduous forests of Hokkaido, the results suggest the necessity to consider the whole list of identified terrestrial pollen taxa for generating robust vegetation reconstructions for northern Japan. Applied to the fossil pollen data, both biome-reconstruction approaches demonstrate consistently that oak-dominated cool mixed forest spread in the study region between 5.5 and 3.6 cal ka BP and was subsequently replaced by beech-dominated temperate deciduous forest. The pollen-based climate reconstruction suggests this change in the vegetation composition was caused by a shift from cooler and drier than present climate to warmer and wetter, similar to modern conditions about 3.6 cal ka BP. Comparing the pollen-based reconstruction results with the published marine records from the NW Pacific, the reconstructed vegetation and climate dynamics can be satisfactorily explained by the greater role played by the warm Tsushima Current in the Sea of Japan and in the Tsugaru Strait during the middle and late Holocene. An increase in sea surface temperatures west and south of the study site would favour air temperature rise and moisture uptake and cause an increase in precipitation and snow accumulation in the western part of Hokkaido during the late Holocene.

Temperature tolerance of western Baltic Sea Fucus vesiculosus - growth, photosynthesis and survival

Fucus vesiculosus L. (Phaeophyceae) is the most abundant and hence ecologically most important primary producer, carbon sink and habitat provider in the western Baltic Sea. All F. vesiculosus L. specimens were collected on 23 April 2014 from a depth of 0.2-1 m in the non-tidal Kiel Fjord, western Baltic Sea (54°27'N; 10°12'E), where this species forms dense and almost monospecific stands on stones. After sampling the algal thalli were stored in a refrigerator box with water from the sampling site, transported to Bremerhaven and stored at 10 °C for one day in filtered seawater. Experiments were conducted with vegetative apical tips (6.7±0.5 cm length), the actively growing region of F. vesiculosus, which were randomly selected and cut from 144 different individuals prior to the experiments. These tips were acclimated to laboratory conditions for three days in filtered seawater at 10 °C before the start of the experiment. Furthermore, 30 additional vegetative apices were freeze-dried to document the initial biochemical status of F. vesiculosus in its native habitat. A temperature gradient was installed in a walk-in constant cooling chamber (15 °C) in nine water baths (5, 10, 15, 20, 24, 26, 27, 28 and 29 °C ± 0.1 °C) which were tempered by thermostats (5, 10 and 15 °C: Huber Variostat CC + Pilot ONE, Peter Huber Kältemaschinen GmbH, Offenburg, Germany; 20 and 28 °C: Haake DC3, Thermo Fisher Scientific Inc., Waltham, USA; 24, 26, 27 and 29 °C: Haake DC10). Every temperature treatment consisted of four 2 L glass beakers (n = 4). In each beaker four F. vesiculosus apices were grown in 2 µm-filtered North Sea water diluted with demineralized water in a ratio of 1:1 and enriched with nutrients after Provasoli (1968; 1/10 enrichment), leading to a salinity of about 15.6 which equaled habitat conditions. The algae were exposed to an irradiance of 130 µmol photons m-2 s-1 ±10 % (Powerstar HGI-TS 150 W, OSRAM GmbH, Bad Homburg, Germany) measured at the top of the beaker under a 16:8 h L:D cycle. The media in the beakers was changed every third or fourth day and aerated with artificial air containing 380 ppm CO2 (gas mixing device; HTK Hamburg GmbH, Hamburg, Germany). Before the experiment, the algae were acclimated to the final temperatures in steps of 5 °C for 2 days each, beginning at 10 °C. After 21 days exposure time, three out of four samples per replicate were freeze-dried for further biochemical analyses, and afterwards the thermostats were turned off to reduce the temperature to 16±0.4 °C for another 10 days permitting growth under post-culture conditions.

Sea surface temperature reconstruction based on dinoflagellates from the North Atlantic

The application of quantitative and semiquantitative methods to assemblage data from dinoflagellate cysts shows potential for interpreting past environments, both in terms of paleotemperature estimates and in recognizing water masses and circulation patterns. Estimates of winter sea-surface temperature (WSST) were produced by using the Impagidinium Index (II) method, and by applying a winter-temperature transfer function (TFw). Estimates of summer sea-surface temperature (SSST) were produced by using a summer-temperature transfer function (TFs), two methods based on a temperature-distribution chart (ACT and ACTpo), and a method based on the ratio of gonyaulacoid:protoperidinioid specimens (G:P). WSST estimates from the II and TFw methods are in close agreement except where Impagidinium species are sparse. SSST estimates from TFs are more variable. The value of the G:P ratio for the Pliocene data in this paper is limited by the apparent sparsity of protoperidinioids, which results in monotonous SSST estimates of 14-26°C. The ACT methods show two biases for the Pliocene data set: taxonomic substitution may force 'matches' yielding incorrect temperature estimates, and the method is highly sensitive to the end-points of species distributions. Dinocyst assemblage data were applied to reconstruct Pliocene sea-surface temperatures between 3.5-2.5 Ma from DSDP Hole 552A, and ODP Holes 646B and 642B, which are presently located beneath cold and cool-temperate waters north of 56°N. Our initial results suggest that at 3.0 Ma, WSSTs were a few degrees C warmer than the present and that there was a somewhat reduced north-south temperature gradient. For all three sites, it is likely that SSSTs were also warmer, but by an unknown, perhaps large, amount. Past oceanic circulation in the North Atlantic was probably different from the present.

Ages and tephra layers in Chatham Rise sediment cores

We present sea surface temperature (SST) estimates based on the relative abundances of long-chain C37 alkenones (UK37') in four sediment cores from a transect spanning the subtropical to subantarctic waters across the subtropical front east of New Zealand. SST estimates from UK37' are compared to those derived from foraminiferal assemblages (using the modern analog technique) in two of these cores. Reconstructions of SST in core tops and Holocene sediments agree well with modern average summer temperatures of ~18°C in subtropical waters and ~14°C in subpolar waters, with a 4°-5°C gradient across the front. Down core UK37' SST estimates indicate that the regional summer SST was 4°-5°C cooler during the last glaciation with an SST of ~10°C in subpolar waters and an SST of ~14°C in subtropical waters. Temperature reconstructions from foraminiferal assemblages agree with those derived from alkenones for the Holocene. In subtropical waters, reconstructions also agree with a glacial cooling of 4° to ~14°C. In contrast, reconstructions for subantarctic pre-Holocene waters indicate a cooling of 8°C with glacial age warm season water temperatures of ~6°C. Thus the alkenones suggest the glacial temperature gradient across the front was the same or reduced slightly to 3.5°-4°C, whereas foraminiferal reconstructions suggest it doubled to 8°C. Our results support previous work indicating that the STF remained fixed over the Chatham Rise during the Last Glacial Maximum. However, the differing results from the two techniques require additional explanation. A change in euphotic zone temperature profiles, seasonality of growth, or preferred growth depth must have affected the temperatures recorded by these biologically based proxies. Regardless of the specific reason, a differential response to the environmental changes between the two climate regimes by the organisms on which the estimates are based suggests increased upwelling associated with increased winds and/or a shallowing of the thermocline associated with increased stratification of the surface layer in the last glaciation.

Sea surface temperature reconstruction for Mediterranean Sea samples

Alkenone unsaturation ratios and planktonic delta18O records from sediment cores of the Alboran, Ionian and Levantine basins in the Mediterranean Sea show pronounced variations in paleo-temperatures and -salinities of surface waters over the last 16,000 years. Average sea surface temperatures (SSTs) are low during the last glacial (averages prior to 13,000 years: 11-15°C), vary rapidly at the beginning of the Holocene, and increase to 17-18°C at all sites during S1 formation (dated between 9500 and 6600 calendar years). The modern temperature gradient (2-3°C) between the Mediterranean sub-basins is maintained during formation of sapropel S1 in the Eastern Mediterranean Sea. After S1, SSTs have remained uniform in the Alboran Sea at 18°C and have fluctuated around 20°C in the Ionian and Levantine Basin sites. The delta18O of planktonic foraminifer calcite decreases by 2 per mil from the late glacial to S1 sediments in the Ionian Basin and by 2.8 per mil in the Levantine Basin. In the Alboran Sea, the decrease is 1.7 per mil. Of the 2.8 per mil decrease in the Levantine Basin, the effect of global ice volume accounts for a maximum of 1.05 per mil and the temperature increase explains only a maximum of 1.3 per mil. The remainder is attributed to salinity changes. We use the temperature and salinity estimates to calculate seawater density changes. They indicate that a reversal of water mass circulation is not a likely explanation for increased carbon burial during S1 time. Instead, it appears that intermediate and deep water formation may have shifted to the Ionian Sea approximately 2000 years before onset of S1 deposition, because surface waters were as cold, but saltier than surface water in the Levantine Basin during the Younger Dryas. Sapropel S1 began to form at the same time, when a significant density decrease also occurred in the Ionian Sea.

Palynology and sea surface temperature reconstruction for the mid-Cretaceous Oceanic Anoxic Event 2 at ODP Hole 210-1276A

The mid-Cretaceous is thought to be a greenhouse world with significantly higher atmospheric pCO2 and sea-surface temperatures as well as a much flatter latitudinal thermal gradient compared to the present. This time interval was punctuated by the Cenomanian/Turonian Oceanic Anoxic Event (OAE-2, ~ 93.5 Myr ago), an episode of global, massive organic carbon burial that likely resulted in a large and abrupt pCO2 decline. However, the climatic consequences of this pCO2 drop are yet poorly constrained. We determined the first, high-resolution sea-surface temperature (SST) record across OAE-2 from a deep-marine sedimentary sequence at Ocean Drilling Program (ODP) Site 1276 in the mid-latitudinal Newfoundland Basin, NW Atlantic. By employing the organic palaeothermometer TEX86, we found that SSTs across the OAE-2 interval were extremely high, but were punctuated by a remarkably large cooling (5-11 °C), which is synchronous with the 2.5-5.5 °C cooling in SST records from equatorial Atlantic sites, and the "Plenus Cold Event". Because this global cooling event is concurrent with increased organic carbon burial, it likely acted in response to the associated pCO2 drop. Our findings imply a substantial increase in the latitudinal SST gradient in the proto-North Atlantic during this period of global cooling and reduced atmospheric pCO2, suggesting a strong coupling between pCO2 and latitudinal thermal gradients under greenhouse climate conditions.

Oxygen-isotope composition, temperature and depth of formation of chert at DSDP Leg 62 Holes

We measured oxygen-isotope compositions of 16 siliceous rocks from Deep Sea Drilling Project Sites 463, 464, 465, and 466 (Leg 62). Samples are from deposits that range in age from about 40 to 103 m.y. and that occur at sub-bottom depths of 9 to 461 meters. Mean d18O values range from 28.4 to 36.8 per mil and 36.0 ± 0.3 per mil for quartz-rich and opal-CTrich rocks, respectively. d18O values in chert decrease with increasing sub-bottom depth; the slope of the d18O/depth curve is less steep for Site 464 than for the other sites which indicates that chert at Site 464 formed at higher temperatures than chert at Sites 463, 465, and 466. Temperatures of formation of cherts were 7 to 42°C, using the silica-water fractionation factor of Knauth and Epstein (1976), or 19 to 56°C, using the equation of Clayton et al. (1972). Temperatures in the sediment where the cherts now occur are lower than their isotopically determined temperatures of formation, which means that the cherts record an earlier history when temperatures in the sediment section were greater. Estimated sediment temperatures when the cherts formed are comparable to, but generally slightly lower than, those calculated from Knauth and Epstein's equation. The isotopic composition of cherts is more closely related to environment of formation (diagenetic environment) or paleogeothermal gradients, than to paleoclimates (bottom-water temperatures). Opal-CT-rich rocks may better record paleo-bottom-water temperature. In Leg 62 cherts, better crystallinity of quartz corresponds to lower d18O values; this implies progressively higher temperatures of equilibration between quartz and water during maturation of quartz. The interrelationship of d18O and crystallinity is noted also in continental-margin deposits such as the Monterey Formation - but for higher temperatures. The apparent temperature difference between open-ocean and continental-margin deposits can be explained by the dominant control of temperature on silica transformation in the rapidly deposited continental-margin deposits, whereas time, as well as temperature, has a strong influence on the transformations in open-ocean deposits. Comparisons between the chemistry and d18O values of cherts reveal two apparent trends: both boron and SiO2 increase as d18O increases. However, the correspondence between SiO2 and d18O is only apparent, because the two cherts lowest in SiO2 are also the most deeply buried, so the trend actually reflects depth of burial. The correspondence between boron and d18O supports the conclusion that boron is incorporated in the quartz crystal structure during precipitation

Climate and total tree cover reconstructions based on the 43.7-kyr fossil pollen record from Khoe (NW Sakhalin Island) and modern climate variables of 236 modern surface pollen sampling sites

A late Quaternary pollen record from northern Sakhalin Island (51.34°N, 142.14°E, 15 m a.s.l.) spanning the last 43.7 ka was used to reconstruct regional climate dynamics and vegetation distribution by using the modern analogue technique (MAT). The long-term trends of the reconstructed mean annual temperature (TANN) and precipitation (PANN), and total tree cover are generally in line with key palaeoclimate records from the North Atlantic region and the Asian monsoon domain. TANN largely follows the fluctuations in solar summer insolation at 55°N. During Marine Isotope Stage (MIS) 3, TANN and PANN were on average 0.2 °C and 700 mm, respectively, thus very similar to late Holocene/modern conditions. Full glacial climate deterioration (TANN = -3.3 °C, PANN = 550 mm) was relatively weak as suggested by the MAT-inferred average climate parameters and tree cover densities. However, error ranges of the climate reconstructions during this interval are relatively large and the last glacial environments in northern Sakhalin could be much colder and drier than suggested by the weighted average values. An anti-phase relationship between mean temperature of the coldest (MTCO) and warmest (MTWA) month is documented during the last glacial period, i.e. MIS 2 and 3, suggesting more continental climate due to sea levels that were lower than present. Warmest and wettest climate conditions have prevailed since the end of the last glaciation with an optimum (TANN = 1.5 °C, PANN = 800 mm) in the middle Holocene interval (ca 8.7-5.2 cal. ka BP). This lags behind the solar insolation peak during the early Holocene. We propose that this is due to continuous Holocene sea level transgression and regional influence of the Tsushima Warm Current, which reached maximum intensity during the middle Holocene. Several short-term climate oscillations are suggested by our reconstruction results and correspond to Northern Hemisphere Heinrich and Dansgaard-Oeschger events, the Bølling-Allerød and the Younger Dryas. The most prominent fluctuation is registered during Heinrich 4 event, which is marked by noticeably colder and drier conditions and the spread of herbaceous taxa.

Freshwater mesocosm experiment 2012 in Vancouver, Canada on effects of increased temperature and food chain length on oxygen fluxes and plankton community structure

Over broad thermal gradients, the effect of temperature on aerobic respiration and photosynthesis rates explains variation in community structure and function. Yet for local communities, temperature dependent trophic interactions may dominate effects of warming. We tested the hypothesis that food chain length modifies the temperature-dependence of ecosystem fluxes and community structure. In a multi-generation aquatic food web experiment, increasing temperature strengthened a trophic cascade, altering the effect of temperature on estimated mass-corrected ecosystem fluxes. Compared to consumer-free and 3-level food chains, grazer-algae (2-level) food chains responded most strongly to the temperature gradient. Temperature altered community structure, shifting species composition and reducing zooplankton density and body size. Still, food chain length did not alter the temperature dependence of net ecosystem fluxes. We conclude that locally, food chain length interacts with temperature to modify community structure, but only temperature, not food chain length influenced net ecosystem fluxes.