Showing papers in "Oecologia in 2008"

Wavelet analysis of ecological time series

Abstract: Wavelet analysis is a powerful tool that is already in use throughout science and engineering. The versatility and attractiveness of the wavelet approach lie in its decomposition properties, principally its time-scale localization. It is especially relevant to the analysis of non-stationary systems, i.e., systems with short-lived transient components, like those observed in ecological systems. Here, we review the basic properties of the wavelet approach for time-series analysis from an ecological perspective. Wavelet decomposition offers several advantages that are discussed in this paper and illustrated by appropriate synthetic and ecological examples. Wavelet analysis is notably free from the assumption of stationarity that makes most methods unsuitable for many ecological time series. Wavelet analysis also permits analysis of the relationships between two signals, and it is especially appropriate for following gradual change in forcing by exogenous variables.

Carbon isotopes and water use efficiency: sense and sensitivity

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

Increasing precipitation event size increases aboveground net primary productivity in a semi-arid grassland

Abstract: Water availability is the primary constraint to aboveground net primary productivity (ANPP) in many terrestrial biomes, and it is an ecosystem driver that will be strongly altered by future climate change. Global circulation models predict a shift in precipitation patterns to growing season rainfall events that are larger in size but fewer in number. This “repackaging” of rainfall into large events with long intervening dry intervals could be particularly important in semi-arid grasslands because it is in marked contrast to the frequent but small events that have historically defined this ecosystem. We investigated the effect of more extreme rainfall patterns on ANPP via the use of rainout shelters and paired this experimental manipulation with an investigation of long-term data for ANPP and precipitation. Experimental plots (n = 15) received the long-term (30-year) mean growing season precipitation quantity; however, this amount was distributed as 12, six, or four events applied manually according to seasonal patterns for May–September. The long-term mean (1940–2005) number of rain events in this shortgrass steppe was 14 events, with a minimum of nine events in years of average precipitation. Thus, our experimental treatments pushed this system beyond its recent historical range of variability. Plots receiving fewer, but larger rain events had the highest rates of ANPP (184 ± 38 g m−2), compared to plots receiving more frequent rainfall (105 ± 24 g m−2). ANPP in all experimental plots was greater than long-term mean ANPP for this system (97 g m−2), which may be explained in part by the more even distribution of applied rain events. Soil moisture data indicated that larger events led to greater soil water content and likely permitted moisture penetration to deeper in the soil profile. These results indicate that semi-arid grasslands are capable of responding immediately and substantially to forecast shifts to more extreme precipitation patterns.

Range expansion of a habitat-modifying species leads to loss of taxonomic diversity: a new and impoverished reef state

Abstract: Global climate change is predicted to have major negative impacts on biodiversity, particularly if important habitat-modifying species undergo range shifts. The sea urchin Centrostephanus rodgersii (Diadematidae) has recently undergone poleward range expansion to relatively cool, macroalgal dominated rocky reefs of eastern Tasmania (southeast Australia). As in its historic environment, C. rodgersii in the extended range is now found in association with a simplified 'barrens' habitat grazed free of macroalgae. The new and important role of this habitat-modifier on reef structure and associated biodiversity was clearly demonstrated by completely removing C. rodgersii from incipient barrens patches at an eastern Tasmanian site and monitoring the macroalgal response relative to unmanipulated barrens patches. In barrens patches from which C. rodgersii was removed, there was a rapid proliferation of canopy-forming macroalgae (Ecklonia radiata and Phyllospora comosa), and within 24 months the algal community structure had converged with that of adjacent macroalgal beds where C. rodgersii grazing was absent. A notable scarcity of limpets on C. rodgersii barrens in eastern Tasmania (relative to the historic range) likely promotes rapid macroalgal recovery upon removal of the sea urchin. In the recovered macroalgal habitat, faunal composition redeveloped similar to that from adjacent intact macroalgal beds in terms of total numbers of taxa, total individuals and Shannon diversity. In contrast, the faunal community of the barrens habitat is overwhelmingly impoverished. Of 296 individual floral/faunal taxa recorded, only 72 were present within incipient barrens, 253 were present in the recovered patches, and 221 were present within intact macroalgal beds. Grazing activity of C. rodgersii results in an estimated minimum net loss of approximately 150 taxa typically associated with Tasmanian macroalgal beds in this region. Such a disproportionate effect by a single range-expanding species demonstrates that climate change may lead to unexpectedly large impacts on marine biodiversity as key habitat-modifying species undergo range modification.

Contrasting effects of invasive plants in plant-pollinator networks.

Abstract: The structural organization of mutualism networks, typified by interspecific positive interactions, is important to maintain community diversity However, there is little information available about the effect of introduced species on the structure of such networks We compared uninvaded and invaded ecological communities, to examine how two species of invasive plants with large and showy flowers (Carpobrotus affine acinaciformis and Opuntia stricta) affect the structure of Mediterranean plant-pollinator networks To attribute differences in pollination to the direct presence of the invasive species, areas were surveyed that contained similar native plant species cover, diversity and floral composition, with or without the invaders Both invasive plant species received significantly more pollinator visits than any native species and invaders interacted strongly with pollinators Overall, the pollinator community richness was similar in invaded and uninvaded plots, and only a few generalist pollinators visited invasive species exclusively Invasive plants acted as pollination super generalists The two species studied were visited by 43% and 31% of the total insect taxa in the community, respectively, suggesting they play a central role in the plant-pollinator networks Carpobrotus and Opuntia had contrasting effects on pollinator visitation rates to native plants: Carpobrotus facilitated the visit of pollinators to native species, whereas Opuntia competed for pollinators with native species, increasing the nestedness of the plant-pollinator network These results indicate that the introduction of a new species to a community can have important consequences for the structure of the plant-pollinator network

Leaf nitrogen:phosphorus stoichiometry across Chinese grassland biomes

Abstract: Leaf N and P stoichiometry covaries with many aspects of plant biology, yet the drivers of this trait at biogeographic scales remain uncertain. Recently we reported the patterns of leaf C and N based on systematic census of 213 species over 199 research sites in the grassland biomes of China. With the expanded analysis of leaf P, here we report patterns of leaf P and N:P ratios, and analyze the relative contribution of climatic variables and phylogeny in structuring patterns of leaf N:P stoichiometry. Average values of leaf P and N:P ratio were 1.9 mg g−1 and 15.3 (mass ratio), respectively, consistent with the previous observation of a higher N:P ratio in China’s flora than the global averages (ca. 13.8), resulting from a lower leaf P. Climatic variables had very little direct correlation with leaf P and N:P ratios, with growing season precipitation and temperature together explaining less than 2% of the variation, while inter-site differences and within-site phylogenetic variation explained 55 and 26% of the total variation in leaf P and N:P ratios. Across all sites and species, leaf N and P were highly positively correlated at all levels. However, the within-site, within-species covariations of leaf N and P were weaker than those across sites and across species. Leaf N and P relationships are driven by both variation between sites at the landscape scale (explaining 58% of the variance) and within sites at the local scale (explaining 24%), while the climatic factors exerted limited influence (explaining less than 3%). In addition, leaf N:P ratios in two dominant genera Kobresia and Stipa had different responses to precipitation. This study suggests that geographic variation and between-species variation, rather than climatic variation, are the major determinants of grassland foliar stoichiometry at the biome level.

Do associated microbial abundances impact marine demosponge pumping rates and tissue densities

Abstract: The evolution of marine demosponges has led to two basic life strategies: one involving close associations with large and diverse communities of microorganisms, termed high microbial abundance (HMA) species, and one that is essentially devoid of associated microorganisms, termed low microbial abundance (LMA) species. This dichotomy has previously been suggested to correlate with morphological differences, with HMA species having a denser mesohyl and a more complex aquiferous systems composed of longer and narrower water canals that should necessitate slower seawater filtration rates. We measured mesohyl density for a variety of HMA and LMA sponges in the Florida Keys, and seawater pumping rates for a select group of these sponges using an in situ dye technique. HMA sponges were substantially denser than LMA species, and had per unit volume pumping rates 52-94% slower than the LMA sponges. These density and pumping rate differences suggest that evolutionary differences between HMA and LMA species may have resulted in profound morphological and physiological differences between the two groups. The LMA sponge body plan moves large quantities of water through their porous tissues allowing them to rapidly acquire the small particulate organic matter (POM) that supplies the majority of their nutritional needs. In contrast, the HMA sponge body plan is suited to host large and tightly packed communities of microorganisms and has an aquiferous system that increases contact time between seawater and the sponge/microbial consortium that feeds on POM, dissolved organic matter and the raw inorganic materials for chemolithotrophic sponge symbionts. The two evolutionary patterns represent different, but equally successful patterns and illustrate how associated microorganisms can potentially have substantial effects on host evolution.

Impacts of alien invasive plants on soil nutrients are correlated with initial site conditions in NW Europe

Abstract: Alien invasive plants are capable of modifying ecosystem function However, it is difficult to make generalisations because impacts often appear to be species- and site-specific In this study, we examined the impacts of seven highly invasive plant species in NW Europe (Fallopia japonica, Heracleum mantegazzianum, Impatiens glandulifera, Prunus serotina, Rosa rugosa, Senecio inaequidens, Solidago gigantea) on nutrient pools in the topsoil and the standing biomass We tested if the impacts follow predictable patterns, across species and sites or, alternatively, if they are entirely idiosyncratic To that end, we compared invaded and adjacent uninvaded plots in a total of 36 sites with widely divergent soil chemistry and vegetation composition For all species, invaded plots had increased aboveground biomass and nutrient stocks in standing biomass compared to uninvaded vegetation This suggests that enhanced nutrient uptake may be a key trait of highly invasive plant species The magnitude and direction of the impact on topsoil chemical properties were strongly site-specific A striking finding is that the direction of change in soil properties followed a predictable pattern Thus, strong positive impacts (higher topsoil nutrient concentrations in invaded plots compared to uninvaded ones) were most often found in sites with initially low nutrient concentrations in the topsoil, while negative impacts were generally found under the opposite conditions This pattern was significant for potassium, magnesium, phosphorus, manganese and nitrogen The particular site-specific pattern in the impacts that we observed provides the first evidence that alien invasive species may contribute to a homogenisation of soil conditions in invaded landscapes

Aboveground net primary production dynamics in a northern Chihuahuan Desert ecosystem

Abstract: Aboveground net primary production (ANPP) dynamics are a key element in the understanding of ecosystem processes. For semiarid environments, the pulse-reserve framework links ANPP to variable and unpredictable precipitation events contingent on surficial hydrology, soil moisture dynamics, biodiversity structure, trophic dynamics, and landscape context. Consequently, ANPP may be decoupled periodically from processes such as decomposition and may be subjected to complex feedbacks and thresholds at broader scales. As currently formulated, the pulse-reserve framework may not encompass the breadth of ANPP response to seasonal patterns of precipitation and heat inputs. Accordingly, we examined a 6-year (1999–2004), seasonal record of ANPP with respect to precipitation, soil moisture dynamics, and functional groups in a black grama (Bouteloua eriopoda) grassland and a creosotebush (Larrea tridentata) shrubland in the northern Chihuahuan Desert. Annual ANPP was similar in the grassland (51.1 g/m2) and shrubland (59.2 g/m2) and positively correlated with annual precipitation. ANPP differed among communities with respect to life forms and functional groups and responses to abiotic drivers. In keeping with the pulse-reserve model, ANPP in black grama grassland was dominated by warm-season C4 grasses and subshrubs that responded to large, transient summer storms and associated soil moisture in the upper 30 cm. In contrast, ANPP in creosotebush shrubland occasionally responded to summer moisture, but the predominant pattern was slower, non-pulsed growth of cool-season C3 shrubs during spring, in response to winter soil moisture accumulation and the breaking of cold dormancy. Overall, production in this Chihuahuan Desert ecosystem reflected a mix of warm-temperate arid land pulse dynamics during the summer monsoon and non-pulsed dynamics in spring driven by winter soil moisture accumulation similar to that of cool-temperate regions.

Effectiveness of rodents as local seed dispersers of Holm oaks.

Abstract: In this study we assessed the effectiveness of rodents as dispersers of Quercus ilex in a patchy landscape in southeastern Spain. We experimentally followed the fates of 3,200 marked and weighed acorns from dispersal through the time of seedling emergence over three years. Rodents handled about 99% of acorns, and dispersed 67% and cached 7.4% of the dispersed acorns. Most caches were recovered and consumed, and only 1.3% of the original experimental acorns were found alive in caches the following spring. Dispersal distances were short (mean = 356.2 cm, median = 157 cm) and strongly right-skewed. Heavier acorns were dispersed further and were more likely to be cached and survive than lighter acorns. All caches were in litter or soil, and each contained a single acorn. Rodents moved acorns nonrandomly, mostly to oaks and pines. Most surviving acorns were either in oaks, a poor microhabitat for oak recruitment, or shrubs, a suitable microhabitat for oak recruitment. Our results suggest that rodents, by burying a relatively high proportion of acorns singly in shrubs and pines, act as moderately effective dispersers of Q. ilex. Nonetheless, this dispersal comes at a very heavy cost.

Tracing carbon flow in an arctic marine food web using fatty acid-stable isotope analysis.

Abstract: Global warming and the loss of sea ice threaten to alter patterns of productivity in arctic marine ecosystems because of a likely decline in primary productivity by sea ice algae. Estimates of the contribution of ice algae to total primary production range widely, from just 3 to >50%, and the importance of ice algae to higher trophic levels remains unknown. To help answer this question, we investigated a novel approach to food web studies by combining the two established methods of stable isotope analysis and fatty acid (FA) analysis--we determined the C isotopic composition of individual diatom FA and traced these biomarkers in consumers. Samples were collected near Barrow, Alaska and included ice algae, pelagic phytoplankton, zooplankton, fish, seabirds, pinnipeds and cetaceans. Ice algae and pelagic phytoplankton had distinctive overall FA signatures and clear differences in delta(13)C for two specific diatom FA biomarkers: 16:4n-1 (-24.0+/-2.4 and -30.7+/-0.8 per thousand, respectively) and 20:5n-3 (-18.3+/-2.0 and -26.9+/-0.7 per thousand, respectively). Nearly all delta(13)C values of these two FA in consumers fell between the two stable isotopic end members. A mass balance equation indicated that FA material derived from ice algae, compared to pelagic diatoms, averaged 71% (44-107%) in consumers based on delta(13)C values of 16:4n-1, but only 24% (0-61%) based on 20:5n-3. Our estimates derived from 16:4n-1, which is produced only by diatoms, probably best represented the contribution of ice algae relative to pelagic diatoms. However, many types of algae produce 20:5n-3, so the lower value derived from it likely represented a more realistic estimate of the proportion of ice algae material relative to all other types of phytoplankton. These preliminary results demonstrate the potential value of compound-specific isotope analysis of marine lipids to trace C flow through marine food webs and provide a foundation for future work.

Explaining variation in tropical plant community composition : influence of environmental and spatial data quality

Abstract: The degree to which variation in plant community composition (beta-diversity) is predictable from environmental variation, relative to other spatial processes, is of considerable current interest. We addressed this question in Costa Rican rain forest pteridophytes (1,045 plots, 127 species). We also tested the effect of data quality on the results, which has largely been overlooked in earlier studies. To do so, we compared two alternative spatial models [polynomial vs. principal coordinates of neighbour matrices (PCNM)] and ten alternative environmental models (all available environmental variables vs. four subsets, and including their polynomials vs. not). Of the environmental data types, soil chemistry contributed most to explaining pteridophyte community variation, followed in decreasing order of contribution by topography, soil type and forest structure. Environmentally explained variation increased moderately when polynomials of the environmental variables were included. Spatially explained variation increased substantially when the multi-scale PCNM spatial model was used instead of the traditional, broad-scale polynomial spatial model. The best model combination (PCNM spatial model and full environmental model including polynomials) explained 32% of pteridophyte community variation, after correcting for the number of sampling sites and explanatory variables. Overall evidence for environmental control of beta-diversity was strong, and the main floristic gradients detected were correlated with environmental variation at all scales encompassed by the study (c. 100-2,000 m). Depending on model choice, however, total explained variation differed more than fourfold, and the apparent relative importance of space and environment could be reversed. Therefore, we advocate a broader recognition of the impacts that data quality has on analysis results. A general understanding of the relative contributions of spatial and environmental processes to species distributions and beta-diversity requires that methodological artefacts are separated from real ecological differences.

Effects of growth and tissue type on the kinetics of 13C and 15N incorporation in a rapidly growing ectotherm

Abstract: The use of stable isotopes to investigate animal diets, habitat use, and trophic level requires understanding the rate at which animals incorporate the 13C and 15N from their diets and the factors that determine the magnitude of the difference in isotopic composition between the animal’s diet and that of its tissues. We determined the contribution of growth and catabolic turnover to the rate of 13C and 15N incorporation into several tissues that can be sampled non-invasively (skin, scute, whole blood, red blood cells, and plasma solutes) in two age classes of a rapidly growing ectotherm (loggerhead turtles, Caretta caretta). We found significant differences in C and N incorporation rates and isotopic discrimination factors (Δ13C = δ13Ctissues − δ13Cdiet and Δ15N = δ15Ntissues − δ15Ndiet) among tissues and between age classes. Growth explained from 26 to 100% of the total rate of incorporation in hatchling turtles and from 15 to 52% of the total rate of incorporation in juvenile turtles. Because growth contributed significantly to the rate of isotopic incorporation, variation in rates among tissues was lower than reported in previous studies. The contribution of growth can homogenize the rate of isotopic incorporation and limit the application of stable isotopes to identify dietary changes at contrasting time scales and to determine the timing of diet shifts. The isotopic discrimination factor of nitrogen ranged from −0.64 to 1.77‰ in the turtles’ tissues. These values are lower than the commonly assumed average 3.4‰ discrimination factors reported for whole body and muscle isotopic analyses. The increasing reliance on non-invasive and non-destructive sampling in animal isotopic ecology requires that we recognize and understand why different tissues differ in isotopic discrimination factors.

Effects of sample preparation on stable isotope ratios of carbon and nitrogen in marine invertebrates: implications for food web studies using stable isotopes.

Abstract: Trophic ecology has benefitted from the use of stable isotopes for the last three decades. However, during the last 10 years, there has been a growing awareness of the isotopic biases associated with some pre-analytical procedures that can seriously hamper the interpretation of food webs. We have assessed the extent of such biases by: (1) reviewing the literature on the topic, and (2) compiling C and N isotopic values of marine invertebrates reported in the literature with the associated sample preparation protocols. The factors considered were: acid-washing, distilled water rinsing (DWR), sample type (whole individuals or pieces of soft tissues), lipid content, and gut contents. Two-level ANOVA revealed overall large and highly significant effects of acidification for both δ13C values (up to 0.9‰ decrease) and δ15 N values (up to 2.1‰ decrease in whole individual samples, and up to 1.1‰ increase in tissue samples). DWR showed a weak overall effect with δ13C increments of 0.6‰ (for the entire data set) or decrements of 0.7‰ in δ15 N values (for tissue samples). Gut contents showed no overall significant effect, whereas lipid extraction resulted in the greatest biases in both isotopic signatures (δ13C, up to −2.0‰ in whole individuals; δ15N, up to +4.3‰ in tissue samples). The study analyzed separately the effects of the various factors in different taxonomic groups and revealed a very high diversity in the extent and direction of the effects. Maxillopoda, Gastropoda, and Polychaeta were the classes that showed the largest isotopic shifts associated with sample preparation. Guidelines for the standardization of sample preparation protocols for isotopic analysis are proposed both for large and small marine invertebrates. Broadly, these guidelines recommend: (1) avoiding both acid washing and DWR, and (2) performing lipid extraction and gut evacuation in most cases.

Nutritional quality of gorilla diets: consequences of age, sex, and season

Abstract: We tested the effects of age, sex, and season on the nutritional strategies of a group of mountain gorillas (Gorilla beringei) in the Bwindi Impenetrable National Park, Uganda. Through observations of food intake of individual gorillas and nutritional analyses of dietary components over different seasons and environments, we estimated nutrient intake and evaluated diet adequacy. Our results suggest that the nutritional costs of reproduction and growth affect nutrient intake; growing juveniles and adult females ate more food and more protein per kilogram of metabolic body mass than did silverbacks. The diets of silverback males, adult females, and juveniles contained similar concentrations of protein, fiber, and sugar, indicating that adult females and juveniles did not select higher protein foods than silverbacks but rather consumed more dry matter to ingest more protein. Juveniles consumed more minerals (Ca, P, Mg, K, Fe, Zn, Mn, Mo) per kilogram of body mass than adult females and silverback males, and juveniles consumed diets with higher concentrations of phosphorous, iron, and zinc, indicating that the foods they ate contained higher concentrations of these minerals. Seasonally, the amount of food consumed on a dry weight basis did not vary, but with increased frugivory, dietary concentrations of protein and fiber decreased and those of water-soluble carbohydrates increased. Energy intake did not change over the year. With the exception of sodium, gorillas ate diets that exceeded human nutrient requirements. A better understanding of the relative importance of food quantity and quality for different age-sex classes provides insights into the ways in which gorillas may be limited by food resources when faced with environmental heterogeneity.

Soil nutrients affect spatial patterns of aboveground biomass and emergent tree density in southwestern Borneo

Abstract: Studies on the relationship between soil fertility and aboveground biomass in lowland tropical forests have yielded conflicting results, reporting positive, negative and no effect of soil nutrients on aboveground biomass. Here, we quantify the impact of soil variation on the stand structure of mature Bornean forest throughout a lowland watershed (8–196 m a.s.l.) with uniform climate and heterogeneous soils. Categorical and bivariate methods were used to quantify the effects of (1) parent material differing in nutrient content (alluvium > sedimentary > granite) and (2) 27 soil parameters on tree density, size distribution, basal area and aboveground biomass. Trees ≥10 cm (diameter at breast height, dbh) were enumerated in 30 (0.16 ha) plots (sample area = 4.8 ha). Six soil samples (0–20 cm) per plot were analyzed for physiochemical properties. Aboveground biomass was estimated using allometric equations. Across all plots, stem density averaged 521 ± 13 stems ha−1, basal area 39.6 ± 1.4 m2 ha−1 and aboveground biomass 518 ± 28 Mg ha−1 (mean ± SE). Adjusted forest-wide aboveground biomass to account for apparent overestimation of large tree density (based on 69 0.3-ha transects; sample area = 20.7 ha) was 430 ± 25 Mg ha−1. Stand structure did not vary significantly among substrates, but it did show a clear trend toward larger stature on nutrient-rich alluvium, with a higher density and larger maximum size of emergent trees. Across all plots, surface soil phosphorus (P), potassium, magnesium and percentage sand content were significantly related to stem density and/or aboveground biomass (RPearson = 0.368–0.416). In multiple linear regression, extractable P and percentage sand combined explained 31% of the aboveground biomass variance. Regression analyses on size classes showed that the abundance of emergent trees >120 cm dbh was positively related to soil P and exchangeable bases, whereas trees 60–90 cm dbh were negatively related to these factors. Soil fertility thus had a significant effect on both total aboveground biomass and its distribution among size classes.

Hover flies are efficient pollinators of oilseed rape.

Abstract: Understanding the consequences of declining diversity and abundance of pollinators for crops and floral biodiversity is a major challenge for current conservation ecology. However, most studies on this issue focus on bees, while other invertebrate taxa are largely ignored. We investigated the pollination efficiency of the globally abundant hover fly Episyrphus balteatus on the common crop, oilseed rape (Brassica napus). The study was conducted over a period of 2 consecutive years by means of enclosure experiments at an agricultural site located in Central Hesse (Germany). E. balteatus significantly increased both seed set and yield. This effect was very constant in the 2 years, despite considerable interannual differences in total seed numbers and seed mass. It highlights the important role of hover flies as pollinators of arable crops under varying environmental conditions. In contrast to bees, the effect of E. balteatus was lower at high pollinator densities than at low pollinator densities. This suggests adverse effects of density-dependent factors on pollination efficiency at high densities. Thus, models ignoring the modulating effect of biotic interactions by generally assuming a simple positive relationship between pollinator density and pollination efficiency might not apply to a vital component of the pollinator community.

Foliar δ15N values characterize soil N cycling and reflect nitrate or ammonium preference of plants along a temperate grassland gradient

Abstract: The natural abundance of stable 15N isotopes in soils and plants is potentially a simple tool to assess ecosystem N dynamics. Several open questions remain, however, in particular regarding the mechanisms driving the variability of foliar δ15N values of non-N2 fixing plants within and across ecosystems. The goal of the work presented here was therefore to: (1) characterize the relationship between soil net mineralization and variability of foliar Δδ15N (δ15Nleaf − δ15Nsoil) values from 20 different plant species within and across 18 grassland sites; (2) to determine in situ if a plant’s preference for NO3− or NH4+ uptake explains variability in foliar Δδ15N among different plant species within an ecosystem; and (3) test if variability in foliar Δδ15N among species or functional group is consistent across 18 grassland sites. Δδ15N values of the 20 different plant species were positively related to soil net mineralization rates across the 18 sites. We found that within a site, foliar Δδ15N values increased with the species’ NO3− to NH4+ uptake ratios. Interestingly, the slope of this relationship differed in direction from previously published studies. Finally, the variability in foliar Δδ15N values among species was not consistent across 18 grassland sites but was significantly influenced by N mineralization rates and the abundance of a particular species in a site. Our findings improve the mechanistic understanding of the commonly observed variability in foliar Δδ15N among different plant species. In particular we were able to show that within a site, foliar δ15N values nicely reflect a plant’s N source but that the direction of the relationship between NO3− to NH4+ uptake and foliar Δδ15N values is not universal. Using a large set of data, our study highlights that foliar Δδ15N values are valuable tools to assess plant N uptake patterns and to characterize the soil N cycle across different ecosystems.

Summer kill rates and predation pattern in a wolf-moose system: can we rely on winter estimates?

Abstract: So far the vast majority of studies on large carnivore predation, including kill rates and consumption, have been based on winter studies. Because large carnivores relying on ungulates as prey often show a preference for juveniles, kill rates may be both higher and more variable during the summer season than during the rest of the year leading to serious underestimates of the total annual predation rate. This study is the first to present detailed empirical data on kill rates and prey selection in a wolf-moose system during summer (June-September) as obtained by applying modern Global Positioning System-collar techniques on individual wolves (Canis lupus) in Scandinavia. Moose (Alces alces) was the dominant prey species both by number (74.4%) and biomass (95.6%); 89.9% of all moose killed were juveniles, representing 76.0% of the biomass consumed by wolves. Kill rate in terms of the kilogram biomass/kilogram wolf per day averaged 0.20 (range: 0.07-0.32) among wolf territories and was above, or well above, the daily minimum food requirements in most territories. The average number of days between moose kills across wolf territories and study periods was 1.71 days, but increased with time and size of growing moose calves during summer. Over the entire summer (June-September, 122 days), a group (from two to nine) of wolves killed a total of 66 (confidence interval 95%; 56-81) moose. Incorporation of body growth functions of moose calves and yearlings and wolf pups over the summer period showed that wolves adjusted their kill rate on moose, so the amount of biomass/kilogram wolf was relatively constant or increased. The kill rate was much higher (94-116%) than estimated from the winter period. As a consequence, projecting winter kill rates to obtain annual estimates of predation in similar predator-prey systems may result in a significant underestimation of the total number of prey killed.

Temporal patterns in ambient noise of biological origin from a shallow water temperate reef.

Abstract: A systematic study of the ambient noise in the shallow coastal waters of north-eastern New Zealand shows large temporal variability in acoustic power levels between seasons, moon phase and the time of day. Ambient noise levels were highest during the new moon and the lowest during the full moon. Ambient noise levels were also significantly higher during summer and lower during winter. Bandpass filtering (700–2,000 Hz and 2–15 kHz), combined with snap counts and data from other studies show that the majority of the sound intensity increases could be attributed to two organisms: the sea urchin and the snapping shrimp. The increased intensity of biologically produced sound during dusk, new moon and summer could enhance the biological signature of a reef and transmit it further offshore. Ambient noise generated from the coast, especially reefs, has been implicated as playing a role in guiding pelagic post-larval fish and crustaceans to settlement habitats. Determining a causal link between temporal increases in ambient noise and higher rates of settlement of reef fish and crustaceans would provide support for the importance of ambient underwater sound in guiding the settlement of these organisms.

A trait-based approach to species' roles in stream ecosystems: climate change, community structure, and material cycling.

Abstract: The sustained decline in habitat quality and community integrity highlights the importance of understanding how communities and environmental variation interactively contribute to ecosystem services. We performed a laboratory experiment manipulating effects of acclimation temperature (5, 15, 25, and 35 degrees C) on resource acquisition, assimilation and subsequent ecosystem services provided by eight freshwater mussel species. Our results suggest that although freshwater mussels are broadly categorized as filter feeders, there are distinct nested functional guilds (thermally tolerant and sensitive) associated with their thermal performance. At 35 degrees C, thermally tolerant species have increased resource assimilation and higher rates of contributed ecosystem services (nutrient excretion, benthic-pelagic coupling). Conversely, thermally sensitive species have decreased assimilation rates and display an array of functional responses including increased/decreased benthic-pelagic coupling and nutrient excretion. Although thermally sensitive species may be in poorer physiological condition at warmer temperatures, their physiological responses can have positive effects on ecosystem services. We extrapolated these results to real mussel beds varying in species composition to address how shifts in community composition coupled with climate change may shift their contributed ecological services. Comparative field data indicate that two co-existing, abundant species with opposing thermal performance (Actinonaias ligamentina, Amblema plicata) differentially dominate community biomass. Additionally, communities varying in the relative proportion of these species differentially influence the magnitude (benthic-pelagic coupling) and quality (N:P excretion) of ecosystem services. As species are increasingly threatened by climate change, greater emphasis should be placed on understanding the contribution of physiological stress to the integrity and functioning of ecosystems.

Temporal organization of phytoplankton communities linked to physical forcing

Abstract: The performance of individual phytoplankton species is strongly governed by the thermal stratification's impact on vertical mixing within the water column, which alters the position of phytoplankton relative to nutrients and light. The present study documents shifts in phytoplankton structure and vertical positioning that have accompanied intensified long-term stratification in a natural ecosystem. Ordination analysis is used to extract gradients in phytoplankton composition in Lake Tahoe, an extremely nutrient-poor lake, over a 23-year period of records. Community structure in the 1980s was associated most strongly with resource availability (low nitrogen to phosphorus ratios, deeper euphotic zone depth), while intensified stratification dominated the phytoplankton structure since the late 1990s. Within diatoms, small-sized cells increased with reduced mixing, suggesting that suppressed turbulence provides them with a competitive advantage over large-sized cells. Among the morphologically diverse chlorophytes, filamentous and coenobial forms were favored under intensified stratification. The selection for small-sized diatoms is accompanied by a shoaling trend in their vertical position in the water column. In contrast, the motile flagellates displayed a deeper vertical positioning in recent years, indicating that optimal growth conditions shifted likely due to reduced upwelling of nutrients. As the thermal stratification of lakes and oceans is strongly linked to climate variables, the present study confirms that climate warming will alter phytoplankton structure and dynamics largely through effects on nutrient availability and sinking velocities. Intensified stratification should favor the expansion of small-sized species and species with the capability of buoyancy regulation, which may alter primary productivity, nutrient recycling, and higher trophic productivity.

Escaping an evolutionary trap: preference and performance of a native insect on an exotic invasive host.

Abstract: Exotic plants may act as population sinks or evolutionary traps for native herbivores. The native butterfly Pieris oleracea lays eggs on garlic mustard, Alliaria petiolata, but larvae develop very poorly on this exotic invasive plant. We examined oviposition preference of individual females and larval performance of their offspring for individuals from one area where garlic mustard is well established and one where it is absent. These data were used to assess whether garlic mustard is being incorporated into or excluded from the diet. Females from the area without garlic mustard showed a wide range of preference, families had low larval survival on garlic mustard, and larval survivorship showed no correlation with mothers' preferences. Females from the area with garlic mustard preferred it to the native host, and larval survivorship on garlic mustard was positively correlated with the mother's preference. Individuals surviving on garlic mustard took longer to pupate and weighed >30% less compared to pupae reared on normal hosts. Our results suggest that where garlic mustard is well established P. oleracea may be adapting to this plant by both improved larval performance and increased adult female oviposition preference for it.

Coordination of leaf and stem water transport properties in tropical forest trees

Abstract: Stomatal regulation of transpiration constrains leaf water potential (ΨL) within species-specific ranges that presumably avoid excessive tension and embolism in the stem xylem upstream. However, the hydraulic resistance of leaves can be highly variable over short time scales, uncoupling tension in the xylem of leaves from that in the stems to which they are attached. We evaluated a suite of leaf and stem functional traits governing water relations in individuals of 11 lowland tropical forest tree species to determine the manner in which the traits were coordinated with stem xylem vulnerability to embolism. Stomatal regulation of ΨL was associated with minimum values of water potential in branches (Ψbr) whose functional significance was similar across species. Minimum values of Ψbr coincided with the bulk sapwood tissue osmotic potential at zero turgor derived from pressure–volume curves and with the transition from a linear to exponential increase in xylem embolism with increasing sapwood water deficits. Branch xylem pressure corresponding to 50% loss of hydraulic conductivity (P50) declined linearly with daily minimum Ψbr in a manner that caused the difference between Ψbr and P50 to increase from 0.4 MPa in the species with the least negative Ψbr to 1.2 MPa in the species with the most negative Ψbr. Both branch P50 and minimum Ψbr increased linearly with sapwood capacitance (C) such that the difference between Ψbr and P50, an estimate of the safety margin for avoiding runaway embolism, decreased with increasing sapwood C. The results implied a trade-off between maximizing water transport and minimizing the risk of xylem embolism, suggesting a prominent role for the buffering effect of C in preserving the integrity of xylem water transport. At the whole-tree level, discharge and recharge of internal C appeared to generate variations in apparent leaf-specific conductance to which stomata respond dynamically.

Complementarity effects through dietary mixing enhance the performance of a generalist insect herbivore

Abstract: The ontogenetic niche concept predicts that resource use depends on an organism’s developmental stage. This concept has been investigated primarily in animals that show differing resource use strategies as juveniles and as adults, such as amphibians. We studied resource use and performance in the grasshopper Chorthippus parallelus (Orthoptera, Acrididae) provided with food plant mixtures of either one, three or eight plant species throughout their development. C. parallelus survival and fecundity was highest in the food plant mixture with eight plant species and lowest in the treatments where only one single plant species was offered as food. C. parallelus’ consumption throughout its ontogeny depended on sex, and feeding on different plant species was dependent on a grasshopper’s developmental stage. To depict grasshopper foraging in food plant mixtures compared to foraging on single plant species, we introduce the term “relative forage total” (RFT) based on an approach used in biodiversity research by Loreau and Hector (Nature 413:548–274, 2001). RFT of grasshoppers in food plant mixtures was always higher than what would have been expected from foraging in monocultures. The increase in food consumption was due to an overall increase in feeding on plant species in mixtures compared to consumption of the same species offered as a single diet. Thus we argue that grasshopper foraging exhibits complementarity effects. Our results reinforce the necessity to consider development-related changes in insect herbivore feeding. Thorough information on the feeding ontogeny of insect herbivores could not only elucidate their nutritional ecology but also help to shed light on their functional role in plant communities.

Temporal dynamics of the carbon isotope composition in a Pinus sylvestris stand: from newly assimilated organic carbon to respired carbon dioxide

Abstract: The 13C isotopic signature (C stable isotope ratio; δ13C) of CO2 respired from forest ecosystems and their particular compartments are known to be influenced by temporal changes in environmental conditions affecting C isotope fractionation during photosynthesis. Whereas most studies have assessed temporal variation in δ13C of ecosystem-respired CO2 on a day-to-day scale, not much information is available on its diel dynamics. We investigated environmental and physiological controls over potential temporal changes in δ13C of respired CO2 by following the short-term dynamics of the 13C signature from newly assimilated organic matter pools in the needles, via phloem-transported organic matter in twigs and trunks, to trunk-, soil- and ecosystem-respired CO2. We found a strong 24-h periodicity in δ13C of organic matter in leaf and twig phloem sap, which was strongly dampened as carbohydrates were transported down the trunk. Periodicity reappeared in the δ13C of trunk-respired CO2, which seemed to originate from apparent respiratory fractionation rather than from changes in δ13C of the organic substrate. The diel patterns of δ13C in soil-respired CO2 are partly explained by soil temperature and moisture and are probably due to changes in the relative contribution of heterotrophic and autotrophic CO2 fluxes to total soil efflux in response to environmental conditions. Our study shows that direct relations between δ13C of recent assimilates and respired CO2 may not be present on a diel time scale, and other factors lead to short-term variations in δ13C of ecosystem-emitted CO2. On the one hand, these variations complicate ecosystem CO2 flux partitioning, but on the other hand they provide new insights into metabolic processes underlying respiratory CO2 emission.

Stem and leaf hydraulics of congeneric tree species from adjacent tropical savanna and forest ecosystems

Abstract: Leaf and stem functional traits related to plant water relations were studied for six congeneric species pairs, each composed of one tree species typical of savanna habitats and another typical of adjacent forest habitats, to determine whether there were intrinsic differences in plant hydraulics between these two functional types. Only individuals growing in savanna habitats were studied. Most stem traits, including wood density, the xylem water potential at 50% loss of hydraulic conductivity, sapwood area specific conductivity, and leaf area specific conductivity did not differ significantly between savanna and forest species. However, maximum leaf hydraulic conductance (Kleaf) and leaf capacitance tended to be higher in savanna species. Predawn leaf water potential and leaf mass per area were also higher in savanna species in all congeneric pairs. Hydraulic vulnerability curves of stems and leaves indicated that leaves were more vulnerable to drought-induced cavitation than terminal branches regardless of genus. The midday Kleaf values estimated from leaf vul- nerability curves were very low implying that daily embolism repair may occur in leaves. An electric circuit analog model predicted that, compared to forest species, savanna species took longer for their leaf water potentials to drop from pre- dawn values to values corresponding to 50% loss of Kleaf or to the turgor loss points, suggesting that savanna species were more buffered from changes in leaf water potential. The results of this study suggest that the relative success of savanna over forest species in savanna is related in part to their ability to cope with drought, which is determined more by leaf than by stem hydraulic traits. Variation among genera accounted for a large proportion of the total variance in most traits, which indicates that, despite different selective pres- sures in savanna and forest habitats, phylogeny has a stronger effect than habitat in determining most hydraulic traits.

Salinity mediates the competitive interactions between invasive mosquitofish and an endangered fish.

Abstract: The interplay of abiotic factors and competition has a long history in ecology, although there are very few studies on the interaction of salinity and competition in fish. Mosquitofish (Gambusia holbrooki) are among the most invasive fish worldwide, with well documented ecological impacts on several taxa such as amphibians and small native fish. It has been previously hypothesized, based on field observations, that salinity limits the invasive success of mosquitofish and provides a competitive refuge for Mediterranean cyprinodonts. We experimentally tested this hypothesis by examining the agonistic behaviour and food competition between mosquitofish and an endangered native cyprinodont (Aphanius fasciatus) at three salinities (0, 15, 25 per thousand). Intraspecific aggressive behaviour for both species was not significantly affected by salinity. As salinity increased, mosquitofish decreased their aggressive behaviour towards cyprinodonts and captured less prey. In contrast, the cyprinodonts did not change their behaviour with different salinity treatments, with the possible exception of increased defensive acts in higher salinities, but captured more prey with increasing salinity because of the reduced efficiency of mosquitofish. Our study confirms previous field observations that salinity limits the invasive success of mosquitofish and provides one of the few experimental demonstrations that it may mediate behavioural and competitive interactions between fish species. Condition-specific competition of mosquitofish might be expected with other species and ecosystems worldwide and illustrates the importance of integrating biotic and abiotic factors in the study of interspecific interactions.

A comparative test of phylogenetic diversity indices

Abstract: Traditional measures of biodiversity, such as species richness, usually treat species as being equal. As this is obviously not the case, measuring diversity in terms of features accumulated over evolutionary history provides additional value to theoretical and applied ecology. Several phylogenetic diversity indices exist, but their behaviour has not yet been tested in a comparative framework. We provide a test of ten commonly used phylogenetic diversity indices based on 40 simulated phylogenies of varying topology. We restrict our analysis to a topological fully resolved tree without information on branch lengths and species lists with presence–absence data. A total of 38,000 artificial communities varying in species richness covering 5–95% of the phylogenies were created by random resampling. The indices were evaluated based on their ability to meet a priori defined requirements. No index meets all requirements, but three indices turned out to be more suitable than others under particular conditions. Average taxonomic distinctness (AvTD) and intensive quadratic entropy (J) are calculated by averaging and are, therefore, unbiased by species richness while reflecting phylogeny per se well. However, averaging leads to the violation of set monotonicity, which requires that species extinction cannot increase the index. Total taxonomic distinctness (TTD) sums up distinctiveness values for particular species across the community. It is therefore strongly linked to species richness and reflects phylogeny per se weakly but satisfies set monotonicity. We suggest that AvTD and J are best applied to studies that compare spatially or temporally rather independent communities that potentially vary strongly in their phylogenetic composition—i.e. where set monotonicity is a more negligible issue, but independence of species richness is desired. In contrast, we suggest that TTD be used in studies that compare rather interdependent communities where changes occur more gradually by species extinction or introduction. Calculating AvTD or TTD, depending on the research question, in addition to species richness is strongly recommended.

The invasive species Alliaria petiolata (garlic mustard) increases soil nutrient availability in northern hardwood-conifer forests.

Abstract: The invasion of non-native plants can alter the diversity and activity of soil microorganisms and nutrient cycling within forests. We used field studies to analyze the impact of a successful invasive groundcover, Alliaria petiolata, on fungal diversity, soil nutrient availability, and pH in five northeastern US forests. We also used laboratory and greenhouse experiments to test three mechanisms by which A. petiolata may alter soil processes: (1) the release of volatile, cyanogenic glucosides from plant tissue; (2) the exudation of plant secondary compounds from roots; and (3) the decomposition of litter. Fungal community composition was significantly different between invaded and uninvaded soils at one site. Compared to uninvaded plots, plots invaded by A. petiolata were consistently and significantly higher in N, P, Ca and Mg availability, and soil pH. In the laboratory, the release of volatile compounds from the leaves of A. petiolata did not significantly alter soil N availability. Similarly, in the greenhouse, the colonization of native soils by A. petiolata roots did not alter soil nutrient cycling, implying that the exudation of secondary compounds has little effect on soil processes. In a leaf litter decomposition experiment, however, green rosette leaves of A. petiolata significantly increased the rate of decomposition of native tree species. The accelerated decomposition of leaf litter from native trees in the presence of A. petiolata rosette leaves shows that the death of these high-nutrient-content leaves stimulates decomposition to a greater extent than any negative effect that secondary compounds may have on the activity of the microbes decomposing the native litter. The results presented here, integrated with recent related studies, suggest that this invasive plant may change soil nutrient availability in such a way as to create a positive feedback between site occupancy and continued proliferation.