Hagfish in the New Zealand fjords are supported by chemoautotrophy of forest carbon

Forest litter is often considered to be a minor energy source to marine communities due to its refractory nature. Large volumes of forest litter are deposited in the New Zealand fjords, and likely recycled into available energy by microbial activity. In this study we used evidence from stable isotope analyses to test whether recycled carbon from chemoautotrophs was an important contributor to the diet of hagfish (Eptatretus cirrhatus). We then analyzed fatty acid biomarkers from the chemoautotrophic clam Solemya parkinsoni and E. cirrhatus to further discriminate the contribution of marine, terrestrial, and chemoautotrophic sources. Bulk isotopic signatures of E. cirrhatus varied considerably (delta13C, from -29.2 per thousand to -16.7 per thousand; delta15N, from -2.8 per thousand to +15.5 per thousand; delta34S, from -21.7 per thousand to +16.7 per thousand) and indicated that a significant percentage of organic matter (38-51%) originated from chemoautotrophs (delta13C, -31.3 per thousand +/- 0.1 per thousand [mean +/- SE]; delta15N, -5.7 per thousand +/- 0.2 per thousand; delta34S, -32.per thousand +/- 3.8 per thousand). Fatty acid biomarkers were depleted in 13C, particularly cis-vaccenic acid (18:1omega7: delta13C, -39.0 per thousand) indicating specific microbial origins of carbon. A high proportion of forest litter in sediments, coupled with isotopic and fatty acid biomarker results, indicates that terrestrial organic matter is a dominant contributor to this marine benthic system. This study demonstrates a clear linkage between terrestrial and marine ecological processes.     

Fate of nitrogen in riparian forest soils and trees: an 15N tracer study simulating salmon decay

We introduced an 15N-NH4+ tracer to the riparian forest of a salmon-bearing stream (Kennedy Creek, Washington, USA) to quantify the cycling and fate of a late-season pulse of salmon N and, ultimately, mechanisms regulating potential links between salmon abundance and tree growth. The 15N tracer simulated deposition of 7.25 kg of salmon (fresh) to four 50-m2 plots. We added NH4+ (the initial product of salmon carcass decay) and other important nutrients provided by carcasses (P, S, K, Mg, Ca) to soils in late October 2003, coincident with local salmon spawning. We followed the 15N tracer through soil and tree pools for one year. Biological uptake of the 15N tracer occurred quickly: 64% of the 15N tracer was bound in soil microbiota within 14 days, and roots of the dominant riparian tree, western red cedar (Thuja plicata), began to take up 15N tracer within seven days. Root uptake continued through the winter. The 15N tracer content of soil organic matter reached a maximum of approximately 52%, five weeks after the application, and a relative equilibrium of approximately 40% within five months. Six months after the addition, in spring 2004, at least 37% of the 15N tracer was found in tree tissues: approximately 23% in foliage, approximately 11% in roots, and approximately 3% in stems. Within the stems, xylem and phloem sap contained approximately 96% of the tracer N, and approximately 4% was in structural xylem N. After one year, at least 28% of the 15N tracer was still found in trees, and loss from the plots was only approximately 20%. The large portion of tracer N taken up in the fall and reallocated to leaves and stems the following spring provides mechanistic evidence for a one-year-lagged tree-growth response to salmon nutrients. Salmon nutrients have been deposited in the Kennedy Creek system each fall for centuries, but the system shows no evidence of nutrient saturation. Rates of N uptake and retention are a function of site history and disturbance and also may be the result of a legacy effect, in which annual salmon nutrient addition may lead to increased efficiency of nutrient uptake and use.     

Arthropod food web restoration following removal of an invasive wetland plant.

Restoration of habitats impacted by invasive plants is becoming an increasingly important tool in the management of native biodiversity, though most studies do not go beyond monitoring the abundance of particular taxonomic groups, such as the return of native vegetation. Yet, the reestablishment of trophic interactions among organisms in restored habitats is equally important if we are to monitor and understand how ecosystems recover. This study examined whether food web interactions among arthropods (as inferred by abundance of naturally occurring stable isotopes of C [delta13C] and N [delta15N]) were reestablished in the restoration of a coastal Spartina alterniflora salt marsh that had been invaded by Phragmites australis. From patterns of C and N stable isotopes we infer that trophic interactions among arthropods in the native salt marsh habitats are characterized by reliance on the dominant marsh plant Spartina as a basal resource. Herbivores such as delphacid planthoppers and mirid bugs have isotope signatures characteristic of Spartina, and predatory arthropods such as dolicopodid flies and spiders likewise have delta13C and delta15N signatures typical of Spartina-derived resources (approximately -13 per thousand and 10 per thousand, respectively). Stable isotope patterns also suggest that the invasion of Phragmites into salt marshes and displacement of Spartina significantly alter arthropod food web interactions. Arthropods in Phragmites-dominated sites have delta13C isotope values between -18 per thousand and -20 per thousand, suggesting reliance on detritus and/or benthic microalgae as basal resources and not on Phragmites, which has a delta13C approximately -26 per thousand. Since most Phragmites herbivores are either feeding internally or are rare transients from nearby Spartina, these resources do not provide significant prey resources for other arthropod consumers. Rather, predator isotope signatures in the invaded habitats indicate dependence on detritus/algae as basal resources instead of the dominant vegetation. The reestablishment of Spartina after removal of Phragmites, however, not only returned species assemblages typical of reference (uninvaded) Spartina, but stable isotope signatures suggest that the trophic interactions among the arthropods were also similar in reestablished habitats. Specifically, both herbivores and predators showed characteristic Spartina signatures, suggesting the return of the original grazer-based food web structure in the restored habitats.     

Whole-lake dissolved inorganic 13C additions reveal seasonal shifts in zooplankton diet

Sustained whole-lake additions of 13C-enriched dissolved inorganic carbon (DIC), intended to increase experimentally the delta13C of DIC in the epilimnion of a small lake with high dissolved organic carbon (DOC), were made during three seasonal periods (spring, summer, and autumn). Coupled with carbon and nitrogen stable isotope analysis of zooplankton and several of their putative food sources, these additions were used to investigate seasonal changes in the relative contributions of different food sources to zooplankton diet in the lake. Four main potential food sources were considered: phytoplankton, heterotrophic bacteria (HB), methanotrophic bacteria (MOB), and green sulfur bacteria (GSB). Because the number of potential food sources exceeded the number of isotopes analyzed, a computer program (IsoSource) was used to estimate the range of possible contributions of the various food sources. During all three periods the added inorganic 13C quickly increased the epilimnetic DIC delta13C by between 18 per thousand and 21 per thousand above the initial value of approximately -21 per thousand. This 13C enrichment of DIC was rapidly transmitted to the particulate organic matter (POM), which included photosynthetic phytoplankton. In spring and summer, delta13C of both adult and juvenile Daphnia increased by approximately 10 per thousand, indicating that Daphnia utilized autochthonous carbon. However, this 13C labeling of Daphnia was not so obvious during the autumn period, when their delta13C generally decreased. According to the IsoSource model outputs based on both delta13C and delta15N values, Daphnia utilized all four potential food source types during spring, summer, and autumn, but in different proportions. The possible contribution of phytoplankton to Daphnia diet was substantial (25-71%) in all seasons. The possible contributions of the bacterial food sources were more variable. The possible contribution of GSB was minor (0-20%) at all times and negligible in autumn. The possible contribution of HB was higher but very variable. Methanotrophic bacteria always made a significant contribution to Daphnia diet and were likely the single most important food source in autumn. Since both HB and MOB in this high-DOC lake probably depend largely on allochthonous organic carbon, our results highlight the seasonal variability in the potential importance of ecosystem subsidies in lake food webs.     

Using stable isotopes to reveal shifts in prey consumption by generalist predators.

The effectiveness of generalist predators in biological control may be diminished if increased availability of alternative prey causes individual predators to decrease their consumption of crop pests. Farming practices that enhance densities of microbidetritivores in the detrital food web can lead to increased densities of generalist predators that feed on pest species. The ability to predict the net biocontrol impact of increased predator densities depends upon knowing the extent to which individual predators may shift to detrital prey and feed less on crop pests when prey of the detritus-based food web are more abundant. We addressed this question by comparing ratios of stable isotopes of carbon (delta13C) and nitrogen (delta15N) in generalist ground predators and two types of prey (crop pests and microbidetritivores) in replicated 8 x 8 m cucurbit gardens subjected to one of two treatments: a detrital subsidy or no addition of detritus (control). Small sheet-web spiders (Linyphiidae) and small wolf spiders (Lycosidae) had delta13C values similar to those of Collembola in both the detrital and control treatments, indicating that small spiders belong primarily to the detrital food web. In control plots the larger generalist predators had delta13C values similar to those of the major insect pests, consistent with their known effectiveness as biocontrol agents. Adding detritus may have caused delta13C of one species of large wolf spider to shift toward that of the microbi-detritivores, although evidence is equivocal. In contrast, another large wolf spider displayed no shift in delta13C in the detrital treatment. Thus, stable isotopes revealed which generalist predators will likely continue to feed on pest species in the presence of greater densities of alternative prey.     

Using nocturnal cold air drainage flow to monitor ecosystem processes in complex terrain.

This paper presents initial investigations of a new approach to monitor ecosystem processes in complex terrain on large scales. Metabolic processes in mountainous ecosystems are poorly represented in current ecosystem monitoring campaigns because the methods used for monitoring metabolism at the ecosystem scale (e.g., eddy covariance) require flat study sites. Our goal was to investigate the potential for using nocturnal down-valley winds (cold air drainage) for monitoring ecosystem processes in mountainous terrain from two perspectives: measurements of the isotopic composition of ecosystem-respired CO2 (delta13C(ER)) and estimates of fluxes of CO2 transported in the drainage flow. To test if this approach is plausible, we monitored the wind patterns, CO2 concentrations, and the carbon isotopic composition of the air as it exited the base of a young (approximately 40 yr-old) and an old (>450 yr-old) steeply sided Douglas-fir watershed. Nocturnal cold air drainage within these watersheds was strong, deep, and occurred on more than 80% of summer nights. The depth of cold air drainage rapidly increased to tower height or greater when the net radiation at the top of the tower approached zero. The carbon isotope composition of CO2 in the drainage system holds promise as an indicator of variation in basin-scale physiological processes. Although there was little vertical variation in CO2 concentration at any point in time, we found that the range of CO2 concentration over a single evening was sufficient to estimate delta 13C(ER) from Keeling plot analyses. The seasonal variation in delta 13C(ER) followed expected trends: during the summer dry season delta 13C(ER) became less negative (more enriched in 13C), but once rain returned in the fall, delta 13C(ER) decreased. However, we found no correlation between recent weather (e.g., vapor pressure deficit) and delta 13C(ER) either concurrently or with up to a one-week lag. Preliminary estimates suggest that the nocturnal CO2 flux advecting past the 28-m tower is a rather small fraction (<20%) of the watershed-scale respiration. This study demonstrates that monitoring the isotopic composition and CO2 concentration of cold air drainage at the base of a watershed provides a new tool for quantifying ecosystem metabolism in mountainous ecosystems on the basin scale.     

Nitrogen niches revealed through species and functional group removal in a boreal shrub community

Most theories attempting to explain the coexistence of species in local communities make fundamental assumptions regarding whether neighbors exhibit competitive, neutral, or positive resource-use interactions; however, few long-term data from naturally assembled plant communities exist to test these assumptions. We utilized a 13-year experiment consisting of factorial removal of three shrub species (Vaccinium myrtillus, V. vitis-idaea, and Empetrum hermaphroditum) and factorial removal of two functional groups (tree roots and feather mosses) to assess how neighbors affect N acquisition and growth of each of the three shrub species. The removal plots were established on each of 30 lake islands in northern Sweden that form a natural gradient of resource availability. We tested the hypotheses that: (1) the presence of functionally similar neighbors would reduce shrub N acquisition through competition for a shared N resource; (2) the removal of functional groups would affect shrub N acquisition by altering the breadth of their niches; and (3) soil fertility would influence the effects of neighbor removals. We found that the removal of functionally similar neighbors (i.e., other shrub species) usually resulted in higher biomass and biomass N, with the strength of these effects varying strongly with site fertility. Shrub species removals never resulted in altered stable N isotope ratios (delta(15)N), suggesting that the niche breadth of the three shrubs was unaffected by the presence of neighboring shrub species. In the functional group removal experiment, we found positive effects of feather moss removal on V. myrtillus biomass and biomass N, and negative effects on E. hermaphrotium N concentration and V. vitis-idaea biomass and biomass N. Tree root removal also caused a significant shift in foliar delta(15)N of V. myrtillus and altered the delta(15)N, biomass, and biomass N of E. hermaphroditum. Collectively, these results show that the resource acquisition and niche breadth of the three shrub species are often affected by neighbors, and further that both the identity of neighbors and site fertility strongly determine whether these interactions are positive, negative, or neutral. These findings have implications for understanding species coexistence and the reciprocal relationships between productivity and species diversity in this ecosystem.     

Mercury bioaccumulation and trophic transfer in sympatric snapper species from the Gulf of Mexico.

Consumption of marine fish is a major route of toxic methyl mercury (MeHg) exposure to ocean apex predators and human populations. Here we explore the influence of trophic structure on total mercury (Hg) accumulation in red snapper (RS, Lutjanus campechanus) and gray snapper (GS, Lutjanus griseus) from the coastal Louisiana region of the Gulf of Mexico, west of the Mississippi River. The objectives of this investigation were to: (1) determine the effectiveness of the use of offshore recreational fishing charter boats and marinas as sources of fish samples and (2) compare species differences in Hg bioaccumulation, trophic position, and carbon sources. Our data show that length-normalized Hg concentrations (> or = 97% as MeHg in tissue of both species) were 230% greater in GS in comparison to RS collected from the same general area. Stable C and N isotope signatures (delta15N and delta13C) indicate that GS occupy a slightly higher trophic position (approximately 30% of one trophic position higher) on the Gulf food web in comparison to RS and that GS appear to incorporate higher trophic positioned prey, continually and at smaller sizes. Mercury was strongly correlated with combined delta15N and delta13C in pooled species data, arguing that most of the substantial difference in Hg bioaccumulation between RS and GS can be explained by modest differences in their trophic position and, to a lesser degree, carbon sources, which had low variation and high overlap among species. These observations demonstrate that even minor to moderate differences in trophic position and food habits in sympatric species can create relatively large differences in bioaccumulation regimes and underscores the importance of quantitative characterization of trophic structure in marine MeHg bioaccumulation studies.     

Birds transport nutrients to fragmented forests in an urban landscape.

The influence of urbanization on nutrient cycling is vaguely known. Here we document that birds, especially those increasing in urban areas (such as crows, Corvus macrorhynchos and C. corone), affect nutrient cycles. Using fecal traps, we measured phosphorus (P) and nitrogen (N) input from the excrement of birds in fragmented forests in an urban landscape. Sources of avian feces were examined on the basis of carbon (C), N, and P percentages and stable isotopes of delta15N and delta13C. Nitrogen and P input was aggregated in the urban landscape, being especially high at the forest where crows roosted during winter. The annual P input due to bird droppings (range 0.068-0.460 kg x ha(-1) x yr(-1); mean 0.167 kg x ha(-1) x yr(-1)) was 12.4% of the total of other pathways in typical forests and 52.9% in the evergreen forest where crows roosted. The annual N input due to bird droppings (range 0.44-3.49 kg x ha(-1) x yr(-1); mean 1.15 kg x ha(-1) x yr(-1)) was 5.2% of the total of other pathways in typical forests and 27.0% in the evergreen forest used by roosting crows. Expected sources of nutrients in feces included insects in the breeding season, fruits in autumn, and mammals and birds in winter. Stable isotopes suggested that the source of nutrients in forests used by roosting crows was from outside the forest. Therefore, birds played a significant role as transporters of nutrients from garbage (including fish, livestock, and/or C4 plants such as corn, with high delta15N and delta13C) in residential and business areas to fragmented evergreen forests, especially near their winter roosts.     

Recycle of buried macroalgal detritus in sediments: use of dual-labelling experiments in the field

In intertidal sediments, burial and decomposition of macroalgal detritus can fuel the sediment of carbon (C) and nitrogen (N), which can be either promptly mineralised or assimilated to enter the food web. This study investigates the transfer of algal-derived C and N to the sediment and to the infauna feeding primarily on benthic diatoms. Thalli of Ulva spp. were ¹³C- and ¹⁵N-labelled in the mesocosm and frozen to create detritus. Thawed macroalgae were, then, buried in the sediment of an intertidal sand-flat forming a mosaic of small patches (50 × 50 cm²) enriched with the macroalgae interspersed with bare sediment. The area was dominated by Corophium volutator and Hydrobia ulvae. The uptake of ¹³C and ¹⁵N was measured in the residual macroalgae, in the sediment and in those animals. Decomposition of detritus was rapid and after 4 weeks the residual biomass was 3% the amount added. Algal-derived ¹³C and ¹⁵N were moved to the sediment. The total amount of ¹³C and ¹⁵N retained in the sediment after completing the decomposition was 3.4 ± 0.5% ¹³C and 2.7 ± 0.6% ¹⁵N the amount decomposed. During the first 2 weeks more N than C was assimilated (1.7% ¹³C and 13.5% ¹⁵N). During the remaining two weeks, N was released from the sediment, while there was little accumulation of C (+6.4 ± 2.0 % ¹³C and −7.7 ± 3.8% ¹⁵N). At the end of the decomposition, animals were ¹⁵N- and ¹³C-labelled. Considering the total accumulation of label in the sediment, they accounted for 3.5 ± 1.8% ¹³C and 25.8 ± 12.9 % ¹⁵N. Similarly, considering the mass of the heavy isotopes gained (¹³C) or lost (¹⁵N) during the remaining 2 weeks, the animals accounted for 4.7 ± 2.1% of the ¹³C in excess and for 18.6 ± 9.1% of the ¹⁵N loss. The transfer of C and N to the sediment and to the surface deposit-feeders can be a relevant mechanism to remove the excess of detritus from the sediment.     

Increasing plant use of organic nitrogen with elevation is reflected in nitrogen uptake rates and ecosystem delta15N

It is hypothesized that decreasing mean annual temperature and rates of nitrogen (N) cycling causes plants to switch from inorganic to organic forms of N as the primary mode of N nutrition. To test this hypothesis, we conducted field experiments and collected natural-abundance delta15N signatures of foliage, soils, and ectomycorrhizal sporocarps along a steep elevation-climate gradient in the White Mountains, New Hampshire, USA. Here we show that with increasing elevation organic forms of N became the dominant source of N taken up by hardwood and coniferous tree species based on dual-labeled glycine uptake analysis, an important confirmation of an emerging theory for the biogeochemistry of the N cycle. Variation in natural abundance foliar delta15N with elevation was also consistent with increasing organic N uptake, though a simple, mass balance model demonstrated that the uptake of delta15N depleted inorganic N, rather than fractionation upon transfer of N from mycorrhizal fungi, best explains variations in foliar delta15N with elevation.     

Geographic assignment of seabirds to their origin: combining morphologic, genetic, and biogeochemical analyses.

Longline fisheries, oil spills, and offshore wind farms are some of the major threats increasing seabird mortality at sea, but the impact of these threats on specific populations has been difficult to determine so far. We tested the use of molecular markers, morphometric measures, and stable isotope (delta15N and delta13C) and trace element concentrations in the first primary feather (grown at the end of the breeding period) to assign the geographic origin of Calonectris shearwaters. Overall, we sampled birds from three taxa: 13 Mediterranean Cory's Shearwater (Calonectris diomedea diomedea) breeding sites, 10 Atlantic Cory's Shearwater (Calonectris diomedea borealis) breeding sites, and one Cape Verde Shearwater (C. edwardsii) breeding site. Assignment rates were investigated at three spatial scales: breeding colony, breeding archipelago, and taxa levels. Genetic analyses based on the mitochondrial control region (198 birds from 21 breeding colonies) correctly assigned 100% of birds to the three main taxa but failed in detecting geographic structuring at lower scales. Discriminant analyses based on trace elements composition achieved the best rate of correct assignment to colony (77.5%). Body measurements or stable isotopes mainly succeeded in assigning individuals among taxa (87.9% and 89.9%, respectively) but failed at the colony level (27.1% and 38.0%, respectively). Combining all three approaches (morphometrics, isotopes, and trace elements on 186 birds from 15 breeding colonies) substantially improved correct classifications (86.0%, 90.7%, and 100% among colonies, archipelagos, and taxa, respectively). Validations using two independent data sets and jackknife cross-validation confirmed the robustness of the combined approach in the colony assignment (62.5%, 58.8%, and 69.8% for each validation test, respectively). A preliminary application of the discriminant model based on stable isotope delta15N and delta13C values and trace elements (219 birds from 17 breeding sites) showed that 41 Cory's Shearwaters caught by western Mediterranean long-liners came mainly from breeding colonies in Menorca (48.8%), Ibiza (14.6%), and Crete (31.7%). Our findings show that combining analyses of trace elements and stable isotopes on feathers can achieve high rates of correct geographic assignment of birds in the marine environment, opening new prospects for the study of seabird mortality at sea.     

计划烧除攀枝花苏铁林区地面覆盖物对苏铁生长和土壤理化性质的影响

选择攀枝花苏铁林下计划烧除试验后苏铁及其根系周围土壤(0~15 cm)为对象,研究火烧对攀枝花苏铁植株生长、叶片生理和苏铁根系周围土壤理化性质的影响,以期为区域攀枝花苏铁种群恢复和生态环境保护提供理论依据.结果表明:计划烧除区域苏铁成年树的株数及株高没有显著变化,新生叶片数目和幼苗株数相对未烧除区域显著增加,增幅分别达201.66%和317.7%;烧除区域苏铁植株新生叶片叶绿素a、叶绿素b、叶绿素a+b,可溶性糖、蛋白质含量,硝酸还原酶(NR)、谷氨酰胺合成酶(GS)活性,叶片碳(C)、氮(N)、磷(P)、钾(K)含量,C/N、N/P比与对照区域新生叶片相比无显著性差异,只有类胡萝卜素含量显著降低.计划烧除改变了苏铁根围土壤(0~15 cm)的理化性质,表现在烧除后的土壤含水量、pH值及总N、P、K含量显著降低;但是,火烧显著提高了土壤有机碳(TOC)、硝态氮(NO3--N)、铵态氮(NH4+-N)含量,微生物量C、N含量也略有升高;这说明火烧有利于增加苏铁林下土壤养分(N)的有效性,为烧除后苏铁快速恢复生长提供有利条件.     

Climatic/edaphic controls on soil carbon/nitrogen response to shrub encroachment in desert grassland.

The proliferation of woody plants in grasslands over the past 100+ years can alter carbon, nitrogen, and water cycles and influence land surface-atmosphere interactions. Although the majority of organic carbon in these ecosystems resides belowground, there is no consensus on how this change in land cover has affected soil organic carbon (SOC) and total nitrogen (TN) pools. The degree to which duration of woody plant occupation, climate, and edaphic conditions have mediated SOC and TN responses to changes in life-form composition are poorly understood. We addressed these issues at a desert grassland site in Arizona, USA, where the leguminous shrub velvet mesquite (Prosopis velutina) has proliferated along an elevation/precipitation/temperature gradient and on contrasting soil morphologic surfaces. On sandy loam complexes of mid-Holocene origin, mean SOC and TN of soils in the grassland matrix increased approximately 68% and approximately 45%, respectively, with increasing elevation. Soil organic carbon pools were comparable and TN pools were approximately 23% higher in Pleistocene-aged clay loam complexes co-occurring with Holocene-aged soils at the upper elevation/climatic zone. Across the site, belowground resources associated with large Prosopis plants were 21-154% (SOC) and 18-127% (TN) higher than those in the grassy matrix. The variance in SOC and TN pools accounted for by Prosopis stem size (a rough surrogate for time of site occupation) was highest at the low- and mid-elevation sites (69-74%) and lowest at the upper elevation site (32-38%). Soil delta15N values ranged from 5.5 per thousand to 6.7 per thousand across the soil/elevation zones but were comparable in herbaceous and shrub-impacted soils and exhibited a weak relationship with Prosopis basal stem diameter (r2 < 0.1) and TN (r2 < 0.08). The SOC delta13C values decreased linearly with increasing Prosopis basal diameter, suggesting that size and isotopic composition of the SOC pool is a function of time of Prosopis site occupation. Isotopic mixture models indicate that encroachment of C3 woody plants has also promoted SOC additions from C4 plant sources, indicative of long-term herbaceous facilitation. Grassy sites in contrasting soil/elevation combinations, initially highly distinctive in their SOC pool size and delta13C, appear to be converging on similar values following approximately 100 years of woody plant proliferation.     

Quantitative approaches to the analysis of stable isotope food web data

Ecologists use stable isotopes (delta13C, delta15N) to better understand food webs and explore trophic interactions in ecosystems. Traditionally, delta13C vs. delta15N bi-plots have been used to describe food web structure for a single time period or ecosystem. Comparisons of food webs across time and space are increasing, but development of statistical approaches for testing hypotheses regarding food web change has lagged behind. Here we present statistical methodologies for quantitatively comparing stable isotope food web data. We demonstrate the utility of circular statistics and hypothesis tests for quantifying directional food web differences using two case studies: an arthropod salt marsh community across a habitat gradient and a freshwater fish community from Lake Tahoe, USA, over a 120-year time period. We calculated magnitude and mean angle of change (theta) for each species in food web space using mean delta13C and delta15N of each species as the x, y coordinates. In the coastal salt marsh, arthropod consumers exhibited a significant shift toward dependence on Spartina, progressing from a habitat invaded by Phragmites to a restored Spartina habitat. In Lake Tahoe, we found that all species from the freshwater fish community shifted in the same direction in food web space toward more pelagic-based production with the introduction of nonnative Mysis relicta and onset of cultural eutrophication. Using circular statistics to quantitatively analyze stable isotope food web data, we were able to gain significant insight into patterns and changes in food web structure that were not evident from qualitative comparisons. As more ecologists incorporate a food web perspective into ecosystem analysis, these statistical tools can provide a basis for quantifying directional food web differences from standard isotope data.     

An invader differentially affects leaf physiology of two natives across a gradient in diversity

Little is known about how exotics influence the ecophysiology of co-occurring native plants or how invader impact on plant physiology may be mediated by community diversity or resource levels. We measured the effect of the widespread invasive forb spotted knapweed (Centaurea maculosa) on leaf traits (leaf dry matter content, specific leaf area, leaf nitrogen percentage, leaf C:N ratios, and delta13C as a proxy for water use efficiency) of two co-occurring native perennial grassland species, Monarda fistulosa (bee balm) and Koeleria macrantha (Junegrass). The impact of spotted knapweed was assessed across plots that varied in functional diversity and that either experienced ambient rainfall or received supplemental water. Impact was determined by comparing leaf traits between identical knapweed-invaded and noninvaded assemblages. Virtually all M. fistulosa leaf traits were affected by spotted knapweed. Knapweed impact, however, did not scale with its abundance; the impact of knapweed on M. fistulosa was similar across heavily invaded low-diversity assemblages and lightly invaded high-diversity assemblages. In uninvaded assemblages, M. fistulosa delta13C, leaf nitrogen, and C:N ratios were unaffected by native functional group richness, whereas leaf dry matter content significantly increased and specific leaf area significantly decreased across the diversity gradient. The effects of spotted knapweed on K. macrantha were weak; instead native functional group richness strongly affected K. macrantha leaf C:N ratio, delta13C, and specific leaf area, but not leaf dry matter content. Leaf traits for both species changed in response to spotted knapweed or functional richness, and in a manner that may promote slower biomass accumulation and efficient conservation of resources. Taken together, our results show that an invader can alter native plant physiology, but that these effects are not a simple function of how many invaders exist in the community.     

Long-term aboveground and belowground consequences of red wood ant exclusion in boreal forest

Despite their ubiquity, the role of ants in driving ecosystem processes both aboveground and belowground has been seldom explored, except within the nest. During 1995 we established 16 ant exclusion plots of approximately 1.1 x 1.1 m, together with paired control plots, in the understory layer of a boreal forest ecosystem in northern Sweden that supports high densities of the mound-forming ant Formica aquilonia, a red wood ant species of the Formica rufa group. Aboveground and belowground measurements were then made on destructively sampled subplots in 2001 and 2008, i.e., 6 and 13 years after set-up. While ant exclusion had no effect on total understory plant biomass, it did greatly increase the relative contribution of herbaceous species, most likely through preventing ants from removing their seeds. This in turn led to higher quality resources entering the belowground subsystem, which in turn stimulated soil microbial biomass and activity and the rates of loss of mass and carbon (C) and nitrogen (N) from litter in litterbags placed in the plots. This was accompanied by losses of approximately 15% of N and C stored in the humus on a per area basis. Ant exclusion also had some effects on foliar stable isotope ratios for both C and N, most probably as a consequence of greater soil fertility. Further, exclusion of ants had multitrophic effects on a microbe-nematode soil food web with three consumer trophic levels and after six years promoted the bacterial-based relative to the fungal-based energy channel in this food web. Our results point to a major role of red wood ants in determining forest floor vegetation and thereby exerting wide-ranging effects on belowground properties and processes. Given that the boreal forest occupies 11% of the Earth's terrestrial surface and stores more C than any other forest biome, our results suggest that this role of ants could potentially be of widespread significance for biogeochemical nutrient cycling, soil nutrient capital, and sequestration of belowground carbon.     

Differential tree and shrub production in response to fertilization and disturbance by coastal river otters in Alaska

We explored the interacting effects of marine-derived nutrient fertilization and physical disturbance introduced by coastal river otters (Lontra canadensis) on the production and nutrient status of pristine shrub and tree communities in Prince William Sound, Alaska, USA. We compared production of trees and shrubs between latrines and non-latrines, while accounting for otter site selection, by sampling areas on and off sites. Nitrogen stable isotope analysis (delta15N) indicated that dominant tree and shrub species assimilated the marine-derived N excreted by otters. In association with this uptake, tree production increased, but shrub density and nonwoody aboveground shrub production decreased. The reduced shrub production was caused by destruction of ramets, especially blueberry (Vaccinium spp.), through physical disturbance by river otters. False azalea (Menziesia ferruginea) ramets were less sensitive to otter disturbance. Although surviving individual blueberry ramets showed a tendency for increased production per plant, false azalea allocated excess N to storage in leaves rather than growth. We found that plant responses to animal activity vary among species and levels of biological organization (leaf, plant, ecosystem). Such differences should be accounted for when assessing the influence of river otters on the carbon budget of Alaskan coastal forests at the landscape scale.     

Refueling while flying: foraging bats combust food rapidly and directly to power flight

Flying vertebrates, such as bats, face exceptionally high energy costs during active flapping flight. Once airborne, energy turnover may exceed basal metabolic rate by a factor of up to 15. Here, we asked whether fuel that powers flight originates from exogenous (dietary nutrients), endogenous sources (mostly body lipids or glycogen), or a combination of both. Since most insectivorous bats fly continuously over relatively long time periods during foraging, we assumed that slowly mobilized glycogen, although suitable for supporting brief sallying flights, is inadequate to power aerial insect-hunting of bats. We hypothesized that the insect-feeding Noctilio albiventris rapidly mobilizes and combusts nutrients from insects it has just eaten instead of utilizing endogenous lipids. We used the stable carbon isotope ratio in the bats' exhaled breath (delta13C(brth)) to assess the origin of metabolized substrates of resting and flying N. albiventris in two nutritional conditions: fasted and recently fed. The breath of fasted resting bats was depleted in 13C in relation to their insect diet (delta13C(diet)), indicating the combustion of 13C depleted body lipids. In contrast to this, delta13C(brth) of bats that had recently fed closely matched delta13C(diet) in both resting and flying bats, suggesting a quick mobilization of ingested nutrients for metabolism. In contrast to most non-volant mammals, bats have evolved the ability to fuel their high energy expenditure rates through the rapid combustion of exogenous nutrients, enabling them to conquer the nocturnal niche of aerial insectivory.     

Spatial and temporal shifts in stable isotope values of the bottom-dwelling shrimp<Emphasis Type="Italic"> Nauticaris marionis</Emphasis> at the sub-Antarctic archipelago

Spatial and temporal dynamics of carbon and nitrogen stable isotope signatures of the bottom-dwelling caridean shrimp Nauticaris marionis were measured during April and May between 1984 and 2000 in the vicinity of Marion Island (the Prince Edward Islands, Southern Ocean). There was one trophic-level enrichment in bulk ¹⁵N and ¹³C signatures between small (<20 mm long) and large (>20 mm) specimens of N. marionis, suggesting distinct trophic differentiation among major shrimp size groups. Both ¹⁵N and ¹³C values of N. marionis increased with the depth, reflecting changes in their diet. There were no clear temporal trends in bulk ¹⁵N signatures of N. marionis. However, compound-specific ¹⁵N measurements of amino acids indicated that N. marionis from the inter-island realm occupied the trophic level of second order carnivores, while similarly sized shrimps in the near-shore realm were at the trophic level of first order carnivores. Compound-specific measurements also identified a change in the source of inorganic nitrogen at the base of the food web between the inter-island and near-shore realms. In contrast to the bulk ¹⁵N values, a significant shift in bulk ¹³C values of N. marionis was observed between 1984 and more recent years. This temporal change appears to be linked to changes in the overall productivity of the Prince Edward Island inter-island system, which could be linked to global climate change.Communicated by J.P. Thorpe, Port Erin