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  • 1
    Publication Date: 2014-12-18
    Description: Herein, we report on the crystal structures of Nb 2 AlC and TiNbAlC—actual composition (Ti 0.45 ,Nb 0.55 ) 2 AlC—compounds determined from Rietveld analysis of neutron diffraction patterns in the 300–1173 K temperature range. The average linear thermal expansion coefficients of a Nb 2 AlC sample in the a and c directions are, respectively, 7.9(5) × 10 −6 and 7.7(5) × 10 −6  K −1 on one neutron diffractometer and 7.3(3) × 10 −6 and 7.0(2) × 10 −6  K −1 on a second diffractometer. The respective values for the (Ti 0.45 ,Nb 0.55 ) 2 AlC composition—only tested on one diffractometer—are 8.5(3) × 10 −6 and 7.5(5) × 10 −6  K −1 . These values are relatively low compared to other MAX phases. Like other MAX phases, however, the atomic displacement parameters (APDs) show that the Al atoms vibrate with higher amplitudes than the Ti and C atoms, and more along the basal planes than normal to them. When the predictions of the APDs obtained from density functional theory are compared to the experimental results, good quantitative agreement is found for the Al atoms. In case of the Nb and C atoms, the agreement was more qualitative.
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  • 2
    Publication Date: 2014-12-18
    Description: In this study, lithium disilicate (LS 2 ) glass samples with different particle sizes ranging from less than 105 to 850 μm were prepared. These specimens were inserted in a Pt-Rh DSC crucible and heated to 850°C at different rates (ϕ = 0.5–30 K/min) to identify their crystallization peaks. The activation energies for the overall crystallization ( E ) and the Avrami coefficient ( n ) were evaluated using different nonisothermal models. Specifically, n was evaluated using the Augis–Benett model and the Ozawa method, and E was evaluated using the Kissinger and Ligero methods. As expected, the coarse particles mainly crystallized in the volume, while surface crystallization was predominant in the samples with particle sizes of less than 350 μm. This result was confirmed through SEM analysis of the double stage heat-treated samples. In contrast with previous studies, our results demonstrated that the activation energy decreased as the particle size increased. In addition, no clear correlation between the peak intensity (δ T p ) and the particle size was observed.
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  • 3
    Publication Date: 2014-12-16
    Description: In this letter, 2-μm Pb 0.97 La 0.02 (Zr 0.75 Sn 0.18 Ti 0.07 )O 3 antiferroelectric thick film with tetragonal structure was prepared. The effects of operating electric field, temperature, and frequency on the thermal–electrical energy harvesting capacity of the film were studied by using the Olsen cycle. The results demonstrated that giant energy harvesting effect could be realized in the antiferroelectric thick film. The maximum harvestable energy density per cycle of the film was about 7.8 J/cm 3 at 1 kHz, which was the largest reported value to date. The corresponding energy harvesting efficiency was 0.53%. Moreover, the film had a low leakage current density (about 7.3 × 10 −7 and 3.9 × 10 −5  A/cm 2 at 25 and 200°C, respectively), which was favorable for its application in the devices of the thermal–electrical energy harvesting.
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  • 4
    Publication Date: 2014-12-16
    Description: Discharged energy properties of PbO–SrO–Na 2 O–Nb 2 O 5 –SiO 2 glass-ceramics with crystallization time from 1 to 1000 min were investigated by measuring their hysteresis loops (described as quasi-static measuring method) and pulse-discharge current-time curves (described as dynamic measuring method). The results show the same trend for both measuring methods: With the increment of crystallization time, the discharged energy density increases gradually, while the energy efficiency decreases. The highest energy efficiencies were obtained in the sample with crystallization time of 1 min, which are 96.3% and 82.4%, corresponding quasi-static and dynamic measurement, respectively. The reduction of energy efficiency with crystallization time is attributed to combined effect of ferroelectric polarization and interfacial polarization, and part of the corresponding energy could not release in the pulse-discharge process.
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  • 5
    Publication Date: 2014-12-02
    Description: New lead-free perovskite solid solution ceramics of (1  − x )( Bi 1/2 Na 1/2 ) TiO 3 – x Ba ( Ni 1/2 Nb 1/2 ) O 3 [(1− x )BNT– x BNN, x  =   0.02–0.06) were prepared and their dielectric, ferroelectric, piezoelectric, and electromechanical properties were investigated as a function of the BNN content. The X-ray diffraction results indicated that the addition of BNN has induced a morphotropic phase transformation from rhombohedral to pseudocubic symmetry approximately at x  =   0.045, accompanying an evolution of dielectric relaxor behavior as characterized by enhanced dielectric diffuseness and frequency dispersion. In the proximity of the ferroelectric rhombohedral and pseudocubic phase coexistence zone, the x  =   0.045 ceramics exhibited optimal piezoelectric and electromechanical coupling properties of d 33 ~121 pC/N and k p ~0.27 owing to decreased energy barriers for polarization switching. However, further addition of BNN could cause a decrease in freezing temperatures of polar nanoregions till the coexistence of nonergodic and ergodic relaxor phases occurred near room temperature, especially for the x  =   0.05 sample which has negligible negative strains and thus show the maximum electrostrain of 0.3% under an external electric field of 7 kV/mm, but almost vanished piezoelectric properties. This was attributed to the fact that the induced long-range ferroelectric order could reversibly switch back to its original ergodic state upon removal of external electric fields.
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  • 6
    Publication Date: 2014-12-02
    Description: Light transmission in polycrystalline magnesium fluoride was studied as a function of the mean grain size at different wavelengths. The mean grain size was varied by annealing hot-pressed billets in argon atmosphere at temperatures ranging from 600°C to 800°C for 1 h. The grain-size and grain-orientation distributions were characterized by electron back scatter diffraction. The scattering coefficients were calculated from the in-line transmittance measured at various wavelengths. The scattering coefficient of polycrystalline magnesium fluoride increased linearly with the mean grain size and inversely with the square of the wavelength of light. It is shown that these trends are consistent with theoretical models based on both a limiting form of the Raleigh–Gans–Debye (RGD) theory of particle scattering and light retardation theories that take refractive index variations along the light path. Quantitative predictions of the theories are, however, subject to uncertainly due to the restrictive assumptions made in the theories and difficulties in representing the microstructure in the theoretical models. In particular, grain-size distribution has a significant influence on the scattering coefficient calculated using particle scattering models.
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  • 7
    Publication Date: 2014-12-02
    Description: Determining the absolute chronology of ceramic artifacts has significant implications for archeological and historical research. Wilson, Hall et al . recently suggested a new technique for direct absolute dating of archeological ceramics based on a moisture-induced chemical reaction, called rehydroxylation (RHX) dating. RHX dating proceeds by measuring the mass of chemically combined water in the ceramics in the form of OH hydroxyls, and the mass gain rate at the Effective Lifetime Temperature (ELT) that the ceramics experienced over its lifetime. To date, ELT determinations have been based on estimates of the ceramic's lifetime temperature history; taking into account weather and climate data and the depth at which the artifact was found. The uncertainty in determining the ELT can be a major component of the overall dating uncertainty. Here, we propose an alternative method which relies minimally on weather and climate data, and provides more precise determinations of the ELT and the ceramic age . The proposed method (SAS: Same Age Samples) involves a minimum of four measurements of the RHX mass gain rate constant for two ceramic samples of the same age at two temperatures. We show via simulations that the proposed SAS method can determine the ELT with a precision of 0.2 K which is comparable to the best ELT determination based on lifetime temperature history, and also comparable to available microbalance temperature resolutions of around 0.1 K. The corresponding percent age error is then 1.4%, or 43 yr for a 3000-yr-old ceramic. The proposed SAS method should be tested with ceramic samples of different ages, whose ELT are well-known.
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  • 8
    Publication Date: 2014-12-02
    Description: Glasses in the Na 2 O–CaO–SrO–ZnO–SiO 2 system have previously been investigated for suitability as a reagent in Al-free glass polyalkenoate cements (GPCs). These materials have many properties that offer potential in orthopedics. However, their applicability has been limited, to date, because of their poor strength. This study was undertaken with the aim of increasing the mechanical properties of a series of these Zn-based GPC glasses by doping with nitrogen to give overall compositions of: 10Na 2 O–10CaO–20SrO–20ZnO–(40−3 x )SiO 2 – x Si 3 N 4 ( x is the no. of moles of Si 3 N 4 ). The density, glass-transition temperature, hardness, and elastic modulus of each glass were found to increase fairly linearly with nitrogen content. Indentation fracture resistance also increases with nitrogen content according to a power law relationship. These increases are consistent with the incorporation of N into the glass structure in threefold coordination with silicon resulting in extra cross-linking of the glass network. This was confirmed using 29 Si MAS-NMR which showed that an increasing number of Q 2 units and some Q 3 units with extra bridging anions are formed as nitrogen content increases at the expense of Q 1 units. A small proportion of Zn ions are found to be in tetrahedral coordination in the base oxide glass and the proportion of these increases with the presence of nitrogen.
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  • 9
    Publication Date: 2014-01-22
    Description: Ecological Applications, Volume 24, Issue 1, Page 3, January 2014.
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    Topics: Biology
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  • 10
    Publication Date: 2014-01-22
    Description: Ecological Applications, Volume 24, Issue 1, Page 23-24, January 2014.
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  • 11
    Publication Date: 2014-01-22
    Description: Ecological Applications, Volume 24, Issue 1, Page 227-228, January 2014.
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  • 12
    Publication Date: 2014-01-22
    Description: Ecology, Volume 95, Issue 1, Page 68-77, January 2014. Delayed life history effects (DLHEs) occur when fitness in one life stage affects fitness in subsequent life stages. Given their biphasic life cycle, pond-breeding amphibians provide a natural system for studying DLHEs, although these effects are not restricted to species with biphasic life histories. In this study, we used multiple mark–recapture techniques enabled by a large trapping array to monitor components of fitness and resulting DLHEs in a population of the endangered California tiger salamander (Ambystoma californiense). We found that DLHEs are prominent across all life stage transitions and that there is variation in whether selection acts primarily at the individual or cohort level. We also demonstrated that there is more than an order of magnitude variation in mean cohort fitness, providing tremendous variation for DLHEs to act upon. We documented an evolutionary trade-off between mass at emergence and date of emergence, which may play a role in maintaining the variation in mass (fitness) at emergence. A literature review revealed that such high levels of intercohort variation occur in many other pond-breeding amphibians, and that appropriately documenting the magnitude of intercohort variation requires long-term studies (roughly two population turnovers). Given the profound effect that DLHEs can have on population dynamics, quantifying intercohort variation in mean fitness and the level(s) at which selection acts will be very important for developing accurate models of population dynamics. In general, when developing models of population dynamics, more attention should be paid to variation in mean fitness and not just variation in total numbers.
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  • 13
    Publication Date: 2014-01-22
    Description: Ecological Applications, Volume 24, Issue 1, Page 71-83, January 2014. Concerns about bycatch of protected species have become a dominant factor shaping fisheries management. However, efforts to mitigate bycatch are often hindered by a lack of data on the distributions of fishing effort and protected species. One approach to overcoming this problem has been to overlay the distribution of past fishing effort with known locations of protected species, often obtained through satellite telemetry and occurrence data, to identify potential bycatch hotspots. This approach, however, generates static bycatch risk maps, calling into question their ability to forecast into the future, particularly when dealing with spatiotemporally dynamic fisheries and highly migratory bycatch species. In this study, we use boosted regression trees to model the spatiotemporal distribution of fishing effort for two distinct fisheries in the North Pacific Ocean, the albacore (Thunnus alalunga) troll fishery and the California drift gillnet fishery that targets swordfish (Xiphias gladius). Our results suggest that it is possible to accurately predict fishing effort using
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  • 14
    Publication Date: 2014-01-22
    Description: Ecology, Volume 95, Issue 1, Page 30-36, January 2014. Competition plays an important role in structuring the community dynamics of phytophagous insects. As the number and impact of biological invasions increase, it has become increasingly important to determine whether competitive differences exist between native and exotic insects. We conducted a meta-analysis to test the hypothesis that native/exotic status affects the outcome of herbivore competition. Specifically, we used data from 160 published studies to assess plant-mediated competition in phytophagous insects. For each pair of competing herbivores, we determined the native range and coevolutionary history of each herbivore and host plant. Plant-mediated competition occurred frequently, but neither native nor exotic insects were consistently better competitors. Spatial separation reduced competition in native insects but showed little effect on exotics. Temporal separation negatively impacted native insects but did not affect competition in exotics. Insects that coevolved with their host plant were more affected by interspecific competition than herbivores that lacked a coevolutionary history. Insects that have not coevolved with their host plant may be at a competitive advantage if they overcome plant defenses. As native/exotic status does not consistently predict outcomes of competitive interactions, plant–insect coevolutionary history should be considered in studies of competition.
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  • 15
    Publication Date: 2014-01-24
    Description: Ecology, Volume 0, Issue 0, Ahead of Print. Predicting the future trend and viability of populations is an essential task in ecology. Because many populations respond to changing environments, uncertainty surrounding environmental responses must be incorporated into population assessments. However, understanding the effects of environmental variation on population dynamics requires information on several important demographic parameters which are often difficult to estimate. Integrated population models facilitate the integration of time-series data on population size and all existing demographic information from a species, allowing the estimation of demographic parameters for which limited or no empirical data exist. Although these models are ideal for assessments of population viability, they have so far not included environmental uncertainty. We incorporated environmental variation in an integrated population model to account for both demographic and environmental uncertainty in an assessment of population viability. In addition, we used this model to estimate true juvenile survival, an important demographic parameter for population dynamics that is difficult to estimate empirically. We applied this model to assess the past and future population trend of a rare island endemic songbird, the Montserrat Oriole Icterus oberi, which is threatened by volcanic activity. Montserrat Orioles experienced lower survival in years with volcanic ash-fall, causing periodic population declines that were compensated by higher seasonal fecundity in years with high pre-breeding season rainfall. Due to the inclusion of both demographic and environmental uncertainty in the model, the estimated population growth rate in the immediate future was highly imprecise (95% credible interval 0.844 - 1.105), and the probability of extinction after three generations (in the year 2028) was low (2.1%). This projection demonstrates that accounting for both demographic and environmental sources of uncertainty provides a more realistic assessment of the viability of populations under unknown future environmental conditions.
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  • 16
    Publication Date: 2014-01-24
    Description: Ecology, Volume 0, Issue 0, Ahead of Print. Evidence of age-dependent changes in foraging behavior of free-ranging individuals is scarce, especially at older stages. Using the isotopic niche as a proxy of the trophic niche during both the breeding (blood) and inter-nesting (feather) periods, we report here empirical evidence for age-, gender- and breeding status-dependent foraging ecology and examine its potential consequences on subsequent reproduction and survival in an extremely long-lived species, the wandering albatross. Immature wandering albatrosses of both sexes forage in the subtropics (δ13C) and feed at the same trophic position (δ15N) than the adults. In contrast to immature birds, adult females forage on average at northern latitudes than males, with both sexes feeding in the subtropics during the inter-nesting period, and males, not females, favouring subantartic waters during incubation. In contrast to adult females, males showed a unique pattern among birds and mammals of a continuous change with age in their main feeding habitat by foraging progressively further south in colder waters during both the breeding and inter-nesting periods. In males, foraging at higher latitudes (lower feather δ13C values) is associated with a lower probability to breed during the following years compared to other birds, but with no effect on their probability to survive. Foraging in cold and windy waters may be linked to foraging impairment that might explain different life history trade-offs and lower investment in reproduction with age. This key point requires further longitudinal investigations and/or studies examining foraging success and energy budget of birds feeding in different water masses.
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  • 17
    Publication Date: 2014-01-12
    Description: Ecology, Volume 0, Issue 0, Ahead of Print. Interactions between plants and scatterhoarding animals may shift from mutualism to predation as a function of resources available to those animals. Because seed species differ in their nutrient content and defenses to predation, resource selection and cache management by scatterhoarders, and thus seed fate, may also depend on the relative availability of different seed types. We tracked the fates of tagged Castanea dentata, Quercus alba, and Q. rubra seeds presented to rodents in pairwise combinations and found that C. dentata, which has moderate dormancy prior to germination, survived better in the presence of Q. alba (no dormancy) than with Q. rubra (longer dormancy). Decisions made by scatterhoarders in response to the composition of available seed resources can alter the relationship between masting and seed dispersal effectiveness in individual tree species and may have influenced the evolution of asynchrony among species-specific masting patterns in temperate forests. In theory, preferential allocation of certain seed species to storage or consumption may also result in indirect apparent predation by one seed species on another.
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  • 18
    Publication Date: 2014-01-12
    Description: This study presents a thermodynamic analysis to predict the type of initial, amorphous oxide overgrowth (i.e., am - Al 2 O 3 or am - SiO 2 ) on bare Al – Si alloy substrates. This analysis have taken into account the energies associated with both its interfaces (interface between the Al – Si alloy substrate and the thin oxide film and interface between the thin oxide film and vacuum) along with the bulk Gibbs free energy of oxide formation. This developed analysis is then applied for various parameters, such as, Si alloying element content at the substrate/oxide interface, the growth temperature, the oxide film thickness (up to 1 nm), and various low-index crystallographic surfaces of the substrate. It is found that am - SiO 2 overgrowth is thermodynamically preferred for a combination of lower oxide film thickness, lower growth temperature, and lower Si alloying content at the alloy/oxide interface. This is because of the overcompensation of the lower energies of both the interfaces over the bulk Gibbs free energy. Furthermore, it is found that for all cases, am - Al 2 O 3 forms a more stable interface with Al – Si alloy than am - SiO 2.
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  • 19
    Publication Date: 2014-01-22
    Description: Ecological Applications, Volume 24, Issue 1, Page 4-14, January 2014. Although market-based incentives have helped resolve many environmental challenges, conservation markets still play a relatively minor role in wildlife management. Establishing property rights for environmental goods and allowing trade between resource extractors and resource conservationists may offer a path forward in conserving charismatic species like whales, wolves, turtles, and sharks. In this paper, we provide a conceptual model for implementing a conservation market for wildlife and evaluate how such a market could be applied to three case studies for whales (minke [Balaenoptera acutorostrata], bowhead [Balaena mysticetus], and gray [Eschrictius robustus]). We show that, if designed and operated properly, such a market could ensure persistence of imperiled populations, while simultaneously improving the welfare of resource harvesters.
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  • 20
    Publication Date: 2014-01-22
    Description: Ecological Applications, Volume 24, Issue 1, Page 15-23, January 2014. We critique a proposal to use catch shares to manage transboundary wildlife resources with potentially high non-extractive values, and we focus on the case of whales. Because whales are impure public goods, a policy that fails to capture all nonmarket benefits (due to free riding) could lead to a suboptimal outcome. Even if free riding were overcome, whale shares would face four implementation challenges. First, a whale share could legitimize the international trade in whale meat and expand the whale meat market. Second, a legal whale trade creates monitoring and enforcement challenges similar to those of organizations that manage highly migratory species such as tuna. Third, a whale share could create a new political economy of management that changes incentives and increases costs for nongovernmental organizations (NGOs) to achieve the current level of conservation. Fourth, a whale share program creates new logistical challenges for quota definition and allocation regardless of whether the market for whale products expands or contracts. Each of these issues, if left unaddressed, could result in lower overall welfare for society than under the status quo.
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  • 21
    Publication Date: 2014-01-22
    Description: Ecological Applications, Volume 24, Issue 1, Page 142-157, January 2014. Global wetland biodiversity loss continues unabated, driven by increased demand for freshwater. A key strategy for conservation management of freshwater systems is to maintain the quantity and quality of the natural water regimes, including the frequency and timing of flows. Formalizing an ecological model depicting the key ecological components and the underlying processes of cause and effect is required for successful conservation management. Models linking hydrology with ecological responses can prove to be an invaluable tool for robust decision-making of environmental flows. Here, we explored alternative water management strategies and identified maximal strategies for successful long-term management of colonial waterbirds in the Macquarie Marshes, Australia. We modeled fluctuations in breeding abundances of 10 colonial waterbird species over the past quarter century (1986–2010). Clear relationships existed between flows and breeding, both in frequencies and total abundances, with a strong linear relationship for flows 〉200 GL. Thresholds emerged for triggering breeding events in all 10 species, but these varied among species. Three species displayed a sharp threshold response between 100 GL and 250 GL. These had a breeding probability of 0.5 when flows were 〉180 GL and a 0.9 probability of breeding with flows 〉350 GL. The remaining species had a probability greater than 0.5 of breeding with flows 〉400 GL. Using developed models, we examined the effects of five environmental flow management strategies on the variability of flows and subsequent likelihood of breeding. Management to different target volumes of environmental flows affected overall and specific breeding probabilities. The likelihood of breeding for all 10 colonial waterbirds increased from a regulated historical mean (±SD) of 0.36 ± 0.09 to 0.53 ± 0.14, an improvement of 47.5% ± 18.7%. Management of complex ecosystems depends on good understanding of the responses of organisms to the main drivers of change. Considerable opportunity exists for implementing similar frameworks for other ecosystem attributes, following understanding of their responses to the flow regime, achieving a more complete model of the entire ecosystem.
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  • 22
    Publication Date: 2014-01-22
    Description: Ecological Applications, Volume 24, Issue 1, Page 84-93, January 2014. Information on landscape-scale patterns in species distributions and community types is vital for ecological science and effective conservation assessment and planning. However, detailed maps of plant community structure at landscape scales seldom exist due to the inability of field-based inventories to map a sufficient number of individuals over large areas. The Carnegie Airborne Observatory (CAO) collected hyperspectral and lidar data over Kruger National Park, South Africa, and these data were used to remotely identify 〉500 000 tree and shrub crowns over a 144-km2 landscape using stacked support vector machines. Maps of community compositional variation were produced by ordination and clustering, and the importance of hillslope-scale topo-edaphic variation in shaping community structure was evaluated with redundancy analysis. This remote species identification approach revealed spatially complex patterns in woody plant communities throughout the landscape that could not be directly observed using field-based methods alone. We estimated that topo-edaphic variables representing catenal sequences explained 21% of species compositional variation, while we also uncovered important community patterns that were unrelated to catenas, indicating a large role for other soil-related factors in shaping the savanna community. Our results demonstrate the ability of airborne species identification techniques to map biodiversity for the evaluation of ecological controls on community composition over large landscapes.
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  • 23
    Publication Date: 2014-01-22
    Description: Ecology, Volume 95, Issue 1, Page 98-109, January 2014. Grazing, fire, and climate shape mesic grassland communities. With global change altering all three factors, understanding how grasslands respond to changes in these combined drivers may aid in projecting future changes in grassland ecosystems. We manipulated rainfall and simulated grazing (clipping) in two long-term fire experiments in mesic grasslands in North America (NA) and South Africa (SA). Despite their common drivers, grasslands in NA and SA differ in evolutionary history. Therefore, we expected community structure and production in NA and SA to respond differently to fire, grazing, and drought. Specifically, we hypothesized that NA plant community composition and production would be more responsive than the SA plant communities to changes in the drivers and their interactions, and that despite this expected stability of SA grasslands, drought would be the dominant factor controlling production, but grazing would play the primary role in determining community composition at both sites. Contrary to our hypothesis, NA and SA grasslands generally responded similarly to grazing, drought, and fire. Grazing increased diversity, decreased grass cover and production, and decreased belowground biomass at both sites. Drought alone minimally impacted plant community structure, and we saw similar treatment interactions at the two sites. Drought was not the primary driver of grassland productivity, but instead drought effects were similar to or less than grazing and fire. Even though these grasslands differed in evolutionary history, they responded similarly to our fire, grazing, and climate manipulations. Overall, we found community and ecosystem convergence in NA and SA grasslands. Grazing and fire are as important as climate in controlling mesic grassland ecosystems on both continents.
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  • 24
    Publication Date: 2014-01-22
    Description: Ecology, Volume 95, Issue 1, Page 173-184, January 2014. Biodiversity has been shown to increase the temporal stability of community and ecosystem attributes through multiple mechanisms, but these same mechanisms make less consistent predictions about the effects of richness on population stability. The overall effects of biodiversity on population and community stability will therefore depend on the dominant mechanisms that are likely to vary with the nature of biodiversity loss and the degree of environmental variability. We conducted a mesocosm experiment in which we generated a gradient in zooplankton species richness by directly manipulating dominant species and by allowing/preventing immigration from a metacommunity. The mesocosms were maintained under either constant or variable nutrient environments. Population, community, and ecosystem data were collected for five months. We found that zooplankton population and community stability is enhanced in species-rich communities in both constant and variable environments. Species richness increased primarily through the addition of species with low abundance. The communities that were connected to a metacommunity via immigration were the most diverse and the most stable, indicating the importance of both metacommunity dynamics and rare species for stability. We found little evidence for selection effects or overyielding as stabilizing forces. We did find support for asynchronous dynamics and statistical averaging, both of which predict destabilizing effects at the population level. We also found support for weak interactions, which predicts that both populations and communities will become more stable as richness increases. In order to understand the effects of biodiversity loss on stability, we will need to understand when different stabilizing mechanisms tend to operate but also how multiple mechanisms interact.
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  • 25
    Publication Date: 2014-01-23
    Description: Ecological Applications, Volume 0, Issue 0, Ahead of Print. Vegetation-type conversions between grasslands and shrublands have occurred worldwide in semi-arid regions over the last 150 years. Areas once covered by drought deciduous shrubs in Southern California (Coastal Sage Scrub) are converting to grasslands dominated by non-native species. Increasing fire frequency, drought, and nitrogen deposition have all been hypothesized as causes of this conversion, though there is little direct evidence. We constructed rain-out shelters in a Coastal Sage Scrub community following a wildfire, manipulated water and nitrogen input in a split plot design, and collected annual data on community composition for four years. While shrub cover increased through time in all plots during the post-fire succession, both drought and nitrogen significantly slowed recovery. Four years after the fire, average native shrub cover ranged from over 80% in water addition, ambient nitrogen plots to 20% in water reduction, nitrogen addition plots. European grass cover was high following the fire and remained high in the water reduction plots through the third spring after the fire before decreasing in the fourth year of the study. Adding nitrogen decreased the cover of native plants and increased the cover of Eurasian grasses, but also increased growth of crown-sprouting individuals of one shrub species. Our results suggest that extreme drought during post-fire succession may slow or alter succession, possibly facilitating vegetation-type conversion of Coastal Sage Scrub to grassland. Nitrogen addition slowed succession and, when combined with drought, significantly decreased native cover and increased grass cover. Fire, drought, and atmospheric N deposition are widespread aspects of environmental change that occur simultaneously in this system. Our results suggest these drivers of change may reinforce each other, leading to a continued decline of native shrubs and conversion to annual grassland.
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  • 26
    Publication Date: 2014-01-23
    Description: Ecology, Volume 0, Issue 0, Ahead of Print. Theory of invasion ecology indicates that the number of invading individuals (propagule size) and the timing of invasion are important for invasion success. Propagule size affects establishment success due to an Allee effect and the effect of demographic stochasticity, whereas the timing of invasion does so via niche opportunity produced by fluctuating predation pressure and resource abundance. We propose a synthesis of these two mechanisms by a time-varying dose-response curve where the dose is propagule size and the response is establishment probability. We show an example of the synthesis in a simple predator-prey model where successful invasion occurs as a demographic regime shift because of the bistability of the system. The two mechanisms are not independent, but simultaneously determine invasion success in our model. We found that positive growth rate of an invading species does not ensure its establishment, especially when its propagule size is small or when its growth rate is in a decreasing trend. We suggest the difficulty of understanding invasion process based on a dose-response curve of propagule size as no unique curve can be determined due to the effects of invasion timing (i.e., the threshold of demographic regime shift is time-varying). The results of our model analysis also have an implication on the phase relationship between population cycles of predators and prey.
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  • 27
    Publication Date: 2014-01-23
    Description: Ecology, Volume 0, Issue 0, Ahead of Print. Grasslands have been lost and degraded in the USA since Euro-American settlement due to agriculture, development, introduced invasive species, and changes in fire regimes. Fire is frequently used in prairie restoration to control invasion by trees and shrubs, but may have additional consequences. For example, fire might reduce damage by herbivore and pathogen enemies by eliminating litter, which harbors eggs and spores. Less obviously, fire might influence enemy loads differently for native and introduced plant hosts. We used a controlled burn in a Willamette Valley (Oregon) prairie to examine these questions. We expected that without fire, introduced host plants should have less damage than native host plants because the introduced species are likely to have left many of their enemies behind when they were transported to their new range (the enemy release hypothesis, or ERH). If the ERH holds, then fire, which should temporarily reduce enemies on all species, should give an advantage to the natives because they should see greater total reduction in damage by enemies. Prior to the burn, we censused herbivore and pathogen attack on eight plant species (five of non-native origin: Bromus hordaceous, Cynosuros echinatus, Galium divaricatum, Schedonorus arundinaceus (=Festuca arundinacea), Sherardia arvensis; and three natives: Danthonia californica, Epilobium minutum, and Lomatium nudicale). The same plots were monitored for two years post-fire. Prior to the burn, native plants had more kinds of damage and more pathogen damage than introduced plants, consistent with the ERH. Fire reduced pathogen damage relative to the controls more for the native than the introduced species, but the effects on herbivory were negligible. Pathogen attack was correlated with plant reproductive fitness, whereas herbivory was not. These results suggest that fire may be useful for promoting some native plants in prairies due to its negative effects on their pathogens.
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  • 28
    Publication Date: 2014-01-23
    Description: Ecological Applications, Volume 0, Issue 0, Ahead of Print. Accelerating climate change and other cumulative stressors create an urgent need to understand the influence of environmental variation and landscape features on the connectivity and vulnerability of freshwater species. Here, we introduce a novel modeling framework for aquatic systems that integrates spatially-explicit, individual-based, demographic and genetic (demogenetic) assessments with environmental variables. To show its potential utility, we simulated a hypothetical network of 19 migratory riverine populations (e.g., salmonids) using a riverscape connectivity and demogenetic model (CDFISH). We assessed how stream resistance to movement -- a function of water temperature, fluvial distance, and physical barriers -- might influence demogenetic connectivity and hence population vulnerability. We present demographic metrics (abundance, immigration, and change in abundance) and also genetic metrics (diversity, differentiation, and change in differentiation), and combine them into a single vulnerability index for identifying populations at risk of extirpation. We considered four realistic scenarios that illustrate the relative sensitivity of these metrics for early detection of reduced connectivity: (1) maximum resistance due to high water temperatures throughout the network, (2) minimum resistance due to low water temperatures throughout the network, (3) increased resistance at a tributary junction caused by a partial barrier, and (4) complete isolation of a tributary, leaving resident individuals only. We then apply this demogenetic framework using empirical data for a bull trout metapopulation in the upper Flathead River system, Canada and USA, to assess how current and predicted future stream warming may influence population vulnerability. Results suggest that warmer water temperatures and associated barriers to movement (e.g., low flows, de-watering) are predicted to fragment suitable habitat for migratory salmonids, resulting in the loss of genetic diversity and reduced sizes in certain vulnerable population. This demogenetic simulation framework, which is illustrated in a web-based interactive mapping prototype (http://ptolemy.dbs.umt.edu/pvm/), should be useful for evaluating population vulnerability in a wide variety of dendritic and fragmented riverscapes, helping to guide conservation and management efforts for freshwater species.
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  • 29
    Publication Date: 2014-01-23
    Description: Ecological Applications, Volume 0, Issue 0, Ahead of Print. Tree and shrub abundance has increased in many grasslands causing changes in ecosystem carbon and nitrogen pools that are related to patterns of woody plant distribution. However, with regard to spatial patterns of shrub proliferation, little is known about (i) how they are influenced by grazing or (ii) the extent to which they are influenced by intraspecific interactions. We addressed these questions by quantifying changes in the spatial distribution of Prosopis velutina (mesquite) shrubs over 74 years on grazed and protected grasslands. Livestock are effective agents of mesquite dispersal and mesquite plants have lateral roots extending well beyond the canopy. We therefore hypothesized that mesquite distributions would be (a) random on grazed areas mainly due to cattle dispersion and clustered on protected areas due to decreased dispersal and interspecific interference with grasses; and (b) that clustered or random distributions at early stages of encroachment would give way to regular distributions as stands matured and density-dependent interactions intensified. Assessments in 1932, 1948 and 2006 supported the first hypothesis, but we found no support for the second. In fact, clustering intensified with time on the protected area and the pattern remained random on the grazed site. Although shrub density increased on both areas between 1932 and 2006, we saw no progression toward a regular distribution indicative of density-dependent interactions. We propose that processes related to seed dispersal, grass-shrub seedling interactions, and hydrological constraints on shrub size interact to determine vegetation structure in grassland-to-shrubland state changes with implications for ecosystem function and management.
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  • 30
    Publication Date: 2014-01-11
    Description: Ecological Applications, Volume 0, Issue 0, Ahead of Print. Mounting evidence now shows that fishing activity modifies both heritable life-history traits and ecological processes in harvested populations. However, ecological and evolutionary changes are intimately linked and can occur on the same time-scale, and few studies have investigated their combined effect on fish population dynamics. Here, we contrast two population subunits of a harvested fish species in the Northeast Atlantic, the European hake (Merluccius merluccius), in the light of the emerging field of evolutionary demography, which considers the interacting processes between ecology and evolution. The two subunits experienced similar age/size truncation due to size-selective fishing, but displayed differences in key ecological processes (recruitment success) and phenotypic characteristics (maturation schedule). We investigate how temporal variation in maturation and recruitment success interactively shape the population dynamics of the two subunits. We document that the two subunits of European hake displayed different responses to fishing in maturation schedules, possibly because of the different level of adaptive phenotypic plasticity. Our results also suggest that high phenotypic plasticity can dampen the effects of fisheries-induced demographic truncation on population dynamics, whereas a population subunit characterized by low phenotypic plasticity may suffer from additive effects of ecological and life-history responses. Similar fishing pressure may thus trigger contrasting interactions between life history variation and ecological processes within the same population. The presented findings improve our understanding of how fishing impacts eco-evolutionary dynamics, which is a keystone for a more comprehensive management of harvested species.
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  • 31
    Publication Date: 2014-01-11
    Description: Ecological Applications, Volume 0, Issue 0, Ahead of Print. Conservation practitioners, faced with managing multiple threats to biodiversity and limited funding, must prioritize investment in different management actions. From an economic perspective, it is routine practice to invest where the highest rate of return is expected. This return-on-investment (ROI) thinking can also benefit species conservation, and researchers are developing sophisticated approaches to support decision-making for cost-effective conservation. However, applied use of these approaches is limited. Managers may be wary of 'black-box' algorithms or complex methods that are difficult to explain to funding agencies. As an alternative, we demonstrate the use of a basic ROI analysis for determining where to invest in cost-effective management to address threats to species. This method can be applied using basic geographic information system and spread sheet calculations. We illustrate the approach in a management-action prioritization for a biodiverse region of eastern Australia. We use ROI to prioritize management actions for two threats to a suite of threatened species: habitat degradation by cattle grazing and predation by invasive red foxes (Vulpes vulpes). We show how decisions based on cost-effective threat management depend upon how expected benefits to species are defined and how benefits and costs co-vary. By considering a combination of species richness, restricted habitats, species vulnerability, and costs of management actions, small investments can result in greater expected benefit compared with management decisions that consider only species richness. Furthermore, a landscape management strategy that implements multiple actions is more efficient than managing only for one threat or more traditional approaches that don't consider ROI. Our approach provides transparent and logical decision-support for prioritizing different actions intended to abate threats associated with multiple species; it is of use when managers need a justifiable and repeatable approach to investment.
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  • 32
    Publication Date: 2014-01-12
    Description: Ecological Applications, Volume 0, Issue 0, Ahead of Print. Fire is becoming a pervasive driver of environmental change in Amazonia and is expected to intensify, given projected reductions in precipitation and forest cover. Understanding of the influence of post-deforestation land cover change on fires in Amazonia is limited, even though fires in cleared lands constitute a threat for ecosystems, agriculture, and human health. We used MODIS satellite data to map burned areas annually between 2001 and 2010. We then combined these maps with land cover and climate information to understand the influence of land cover change in cleared lands and dry season severity on fire occurrence and spread in a focus area in the Peruvian Amazon. Fire occurrence, quantified as the probability of burning of individual 232m spatial resolution MODIS pixels was modeled as a function of the area of land cover types within each pixel, drought severity, and distance to roads. Fire spread, quantified as the number of pixels burned in 3x3 pixel windows around each focal burned pixel, was modeled as a function of land cover configuration and area, dry season severity, and distance to roads. We found that vegetation regrowth and oil palm expansion are significantly correlated with fire occurrence but that the magnitude and sign of the correlation depend on drought severity, successional stage of regrowing vegetation and oil palm age. Burning probability increased with the area of non-degraded pastures, fallow, and young oil palm and decreased with larger extents of degraded pastures, secondary forests and adult oil palm plantations. Drought severity had the strongest influence on fire occurrence overriding the effectiveness of secondary forests but not of adult plantations to reduce fire occurrence in severely dry years. Overall, irregular and scattered land cover patches reduced fire spread but irregular and dispersed fallows and secondary forests increased fire spread during dry years. Results underscore the importance of land cover management for reducing fire proliferation in this landscape. Incentives for promoting natural regeneration and perennial crops in cleared lands might help reduce fire risk if those areas are protected against burning in early stages of development and during severely dry years.
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  • 33
    Publication Date: 2014-01-12
    Description: Sintered nanoceramics of Pr -doped lanthanum hafnate, La 2 Hf 2 O 7 : Pr , were prepared by means of a high-pressure sintering technique using nanopowders made by Pechini method. Structure, morphology, and spectroscopic properties of the ceramics compared to the starting powder are presented and discussed. Emission and excitation spectra recorded at room temperature as well as at 7 K using synchrotron radiation are presented together with results of luminescence kinetics measurements. In ceramics, at 7 K, the Pr 3+ luminescence from 3 P 0 (blue-green, green, and red region) and 1 D 2 (red) levels is accompanied by a broad-band emission located in the 380–530 nm range of wavelengths, whereas powders gives only the Pr 3+ -related luminescence. Depending on the excitation wavelength, the broad-band emission maximum moves between 430 and 470 nm indicating superposition of at least two components. In sintered nanoceramics, the lifetimes of Pr 3+ emissions from 3 P 0 and 1 D 2 levels were by 10%–20% shorter compared to the powder. The existence of different luminescence centers was proved by the selective emission decays examination. The fast 5 d → 4 f luminescence of Pr 3+ was not observed from either of the two types of La 2 Hf 2 O 7 :Pr materials.
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  • 34
    Publication Date: 2014-01-15
    Description: Ecology, Volume 0, Issue 0, Ahead of Print. Complementary resource use and redundancy of species that fulfill the same ecological role are two mechanisms that can respectively increase and stabilize process rates in ecosystems. For example, predator complementarity and redundancy can determine prey consumption rates and their stability, yet few studies take into account the multiple predator species attacking multiple prey at different rates in natural communities. Thus, it remains unclear whether these biodiversity mechanisms are important determinants of consumption in entire predator-prey assemblages, such that food-web interaction structure determines community-wide consumption and stability. Here, we use empirical quantitative food-webs to study the community-wide effects of functional complementarity and redundancy of consumers (parasitoids) on herbivore control in temperate forests. We find that complementarity in host resource use by parasitoids was a strong predictor of absolute parasitism rates at the community level, and that redundancy in host-use patterns stabilised community-wide parasitism rates in space, but not through time. These effects can potentially explain previous contradictory results from predator diversity research. Phylogenetic diversity (measured using taxonomic distance) did not explain functional complementarity or parasitism rates, so could not serve as a surrogate measure for functional complementarity. Our study shows that known mechanisms underpinning predator diversity effects on both functioning and stability can easily be extended to link food webs to ecosystem functioning.
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  • 35
    Publication Date: 2014-01-16
    Description: Ecology, Volume 0, Issue 0, Ahead of Print. Floral nectar of many plant species is prone to colonization by microbial organisms such as yeasts. Their presence and metabolism of nectar chemical components have the potential to modify a suite of floral traits important for pollinator attraction, including nectar quality and scent. However, studies on the direct and indirect effects of nectar-inhabiting microogranisms on pollinator behavior and plant reproductive success remain rare. To determine their potential to affect pollinator behavior and plant fitness, we experimentally manipulated the common nectar-inhabiting yeast Metschnikowia reukaufii in the nectar of Delphinium nuttallianum, a short-lived montane perennial herb. We detected positive indirect, pollinator-mediated effects of yeasts on male plant fitness measured as pollen donation using powdered fluorescent dyes. However, we detected no direct or indirect effects on components of female fitness. Matching effects on male plant fitness, pollinators responded positively to the presence of yeasts, removing more nectar from flowers treated with M. reukaufii. Our results provide evidence of effects of nectar-inhabiting yeasts on male plant fitness and highlight the importance of microorganisms in mediating plant-pollinator interactions and subsequent plant fitness.
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  • 36
    Publication Date: 2014-01-17
    Description: Ecological Applications, Volume 0, Issue 0, Ahead of Print. There is a growing need for operational biodiversity mapping methods to quantify and to assess the impact of climate change, habitat alteration and human activity on ecosystem composition and function. Here, we present an original method for the estimation of α- and β-diversity of tropical forests based on high-fidelity imaging spectroscopy. We acquired imagery acquired over high-diversity Amazonian tropical forest landscapes in Perú with the Carnegie Airborne Observatory and developed an unsupervised method to estimate the Shannon Index (H') and variations in species composition using Bray-Curtis Dissimilarity (BC) and non-metric multidimensional scaling (NMDS). An extensive field plot network was used for the validation of remotely sensed α- and β-diversity. Airborne maps of H' were highly correlated with field α-diversity estimates (r = 0.86), and BC was estimated with demonstrable accuracy (r = 0.61-0.76). Our findings are the first direct and spatially-explicit remotely sensed estimates of α- and β-diversity of humid tropical forests, paving the way for new applications using airborne and space-based imaging spectroscopy.
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  • 37
    Publication Date: 2014-01-22
    Description: Ecological Applications, Volume 24, Issue 1, Page 229-233, January 2014.
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  • 38
    Publication Date: 2014-01-22
    Description: Ecological Applications, Volume 24, Issue 1, Page 181-195, January 2014. Stable isotopes are valuable tools for partitioning the components contributing to ecological processes of interest, such as animal diets and trophic interactions, plant resource use, ecosystem gas fluxes, streamflow, and many more. Stable isotope data are often analyzed with simple linear mixing (SLM) models to partition the contributions of different sources, but SLM models cannot incorporate a mechanistic understanding of the underlying processes and do not accommodate additional data associated with these processes (e.g., environmental covariates, flux data, gut contents). Thus, SLM models lack predictive ability. We describe a process-based mixing (PBM) model approach for integrating stable isotopes, other data sources, and process models to partition different sources or process components. This is accomplished via a hierarchical Bayesian framework that quantifies multiple sources of uncertainty and enables the incorporation of process models and prior information to help constrain the source-specific proportional contributions, thereby potentially avoiding identifiability issues that plague SLM models applied to “too many” sources. We discuss the application of the PBM model framework to three diverse examples: temporal and spatial partitioning of streamflow, estimation of plant rooting profiles and water uptake profiles (or water sources) with extension to partitioning soil and ecosystem CO2 fluxes, and reconstructing animal diets. These examples illustrate the advantages of the PBM modeling approach, which facilitates incorporation of ecological theory and diverse sources of information into the mixing model framework, thus enabling one to partition key process components across time and space.
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  • 39
    Publication Date: 2014-01-22
    Description: Ecology, Volume 95, Issue 1, Page 3-8, January 2014. Periodical cicadas (Magicicada spp.) mature in 13 or 17 years, the longest development times for any non-diapausing insects. Selection may favor prolonged development since nymphs experience little mortality and individuals taking 17 years have been shown to have greater fecundity than those taking 13 years. Why don't periodical cicadas take even longer to develop? Nymphs feed on root xylem fluid and move little. Ovipositing females prefer fast-growing trees at forest edges. I hypothesized that (1) adults emerging at edges would be heavier than those from forest interiors and (2) habitat changes could limit development time. I collected newly eclosed females that had neither fed as adults nor moved from their site of development. For M. septendecim, females from edges were 4.9% heavier than those from the interior. I assumed that emergence density indicated habitat quality and measured density at eight sites in 1979, 1996, and 2013. Over three generations, variation in densities was great; densities at two sites crashed, and at one site they exploded to 579/m2. Habitat transience may limit development time because only adults can reassess habitats and reposition offspring. In conclusion, cicadas are affected by habitat characteristics, habitats change over 17 years, and cicadas may emerge, mate, and redistribute their offspring to track habitat dynamics.
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  • 40
    Publication Date: 2014-01-22
    Description: Ecology, Volume 95, Issue 1, Page 56-67, January 2014. Predation has been suggested to be especially important in simple food webs and less productive ecosystems such as the arctic tundra, but very few data are available to evaluate this hypothesis. We examined the hypothesis that avian predators could drive the population dynamics of two cyclic lemming species in the Canadian Arctic. A dense and diverse suite of predatory birds, including the Snowy Owl (Bubo scandiacus), the Rough-legged Hawk (Buteo lagopus), and the Long-tailed Jaeger (Stercorarius longicaudus), inhabits the arctic tundra and prey on collared (Dicrostonyx groenlandicus) and brown (Lemmus trimucronatus) lemmings during the snow-free period. We evaluated the predation pressure exerted by these predators by combining their numerical (variation in breeding and fledgling numbers) and functional (variation in diet and daily consumption rates) responses to variations in lemming densities over the 2004–2010 period. Breeding density and number of fledglings produced by the three main avian predators increased sharply without delay in response to increasing lemming densities. The proportion of collared lemmings in the diet of those predators was high at low lemming density (both species) but decreased as lemming density increased. However, we found little evidence that their daily consumption rates vary in relation to changes in lemming density. Total consumption rate by avian predators initially increased more rapidly for collared lemming but eventually leveled off at a much higher value for brown lemmings, the most abundant species at our site. The combined daily predation rate of avian predators exceeded the maximum daily potential growth rates of both lemming species except at the highest recorded densities for brown lemmings. We thus show, for the first time, that predation pressure exerted without delay by avian predators can limit populations of coexisting lemming species during the snow-free period, and thus, that predation could play a role in the cyclic dynamic of these species in the tundra.
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  • 41
    Publication Date: 2014-01-22
    Description: Ecology, Volume 95, Issue 1, Page 238-239, January 2014.
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  • 42
    Publication Date: 2014-01-22
    Description: Ecology, Volume 95, Issue 1, Page 238-239, January 2014.
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  • 43
    Publication Date: 2014-01-23
    Description: We have investigated the electromechanical response of potassium sodium niobate ( K 0.5 Na 0.5 NbO 3 or KNN) thick films. The high-field strain hysteresis loops and weak-field converse piezoelectric d 33 coefficient of the films were measured and compared with those of KNN bulk ceramics under the same electric field conditions. The converse d 33 values of the thick films and bulk ceramics were equal to 82.5 and 138 pm/V, respectively, at 0.4 kV/mm. The fundamental difference between the piezoelectric response of the KNN films and the ceramics was studied in terms of the effective (“clamped”) piezoelectric d 33 coefficient. The reduction in the piezoelectric d 33 coefficient of the KNN films, resulting from the clamping by the substrate, was compared to lead-based ferroelectric thick films, including Pb ( Zr , Ti ) O 3 (PZT) and (1 −  x ) Pb ( Mg 1/3 Nb 2/3 ) O 3 − x PbTiO 3 (PMN-PT). We propose a possible explanation, based on the particular elastic properties of KNN, for the small relative difference observed between the “clamped” and “unclamped” (“bulk”) d 33 of KNN, in comparison with lead-based systems.
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  • 44
    Publication Date: 2014-01-23
    Description: Lead zirconate titanate (PbZr 1 −  x Ti x O 3 , PZT)/epoxy composites with one- dimensional epoxy in PZT matrix (called 3-1 type piezocomposites) have been fabricated by tert-butyl alcohol (TBA)-based directional freeze casting of PZT matrix and afterward infiltration of epoxy. The composites with PZT volume fraction ranging from 0.36 to 0.69 were obtained by adjusting initial solid loading in freeze-casting slurry. The effect of poling voltage on piezoelectric properties of the composites was studied for various volume fraction of PZT phase. With the increasing of PZT volume fraction, relative permittivity (ε r ) increased linearly and piezoelectric coefficient ( d 33 and d 31 ) increased step by step. The resultant composites with 0.57 PZT volume fraction possessed the highest hydrostatic piezoelectric strain coefficient ( d h ) value (184 pC/N), voltage coefficient ( g h ) value (13.6 × 10 −3  V/m Pa), and hydrostatic figure of merit (HFOM) value (2168 × 10 −15  Pa −1 ).
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  • 45
    Publication Date: 2014-01-23
    Description: Multiphase borosilicate glass-ceramics represent one candidate to contain radioactive nuclear waste separated from used nuclear fuel. In this work, the thermophysical properties from room temperature to 1273 K were investigated for four different borosilicate glass-ceramic compositions containing waste loadings from 42 to 60 wt% to determine the sensitivity of these properties to waste loading, as-fabricated microstructure, and potential evolutions in microstructure brought about by temperature transients. The thermal expansion, specific heat capacity, thermal diffusivity, and thermal conductivity are presented. The impact of increasing waste loading is shown to have a small but measurable effect on the thermophysical properties between the four compositions, contrasted to a much greater impact observed when transitioning from predominantly crystalline to amorphous systems. Thermal cycling below 1273 K was not found to measurably impact the thermophysical properties of the compositions investigated here.
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  • 46
    Publication Date: 2014-03-12
    Description: Ecological Applications, Volume 0, Issue 0, Ahead of Print. Grazing represents one of the most common disturbances in drylands worldwide, affecting both ecosystem structure and functioning. Despite the efforts to understand the nature and magnitude of grazing effects on ecosystem components and processes, contrasting results continue to arise. This is particularly remarkable for the biological soil crust (BSC) communities (i.e. cyanobacteria, lichens and bryophytes), which play an important role in soil dynamics. Here we evaluated simultaneously the effect of grazing impact on BSC communities (resistance) and recovery after livestock exclusion (resilience) in a semiarid grassland of Central Mexico. In particular, we examined BSC species distribution, species richness, taxonomical group cover (i.e., cyanobacteria, lichen, bryophyte) and composition along a disturbance gradient with different grazing regimes (low, medium, high impact) and along a recovery gradient with differently-aged livestock exclosures (short-, medium-, long-term exclusion). Differences in grazing impact and time of recovery from grazing both resulted in slight changes in species richness; however, there were pronounced shifts in species composition and group cover. We found we could distinguish four highly diverse and dynamic BSC species groups: 1) species with high resistance and resilience to grazing, 2) species with high resistance but low resilience, 3) species with low resistance but high resilience, and 4) species with low resistance and resilience. While disturbance resulted in a novel diversity configuration, which may profoundly affect ecosystem functioning, we observed that 10 years of disturbance removal did not lead to the ecosystem structure found after 27 years of recovery. These findings are an important contribution to our understanding of BCS dynamics from a species and community perspective placed in a land use change context.
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  • 47
    Publication Date: 2014-03-13
    Description: Ecology, Volume 0, Issue 0, Ahead of Print. Successful dispersal can enhance both individual fitness and population persistence, but the process of dispersal is often inherently risky. The interplay between the costs and benefits of dispersal are poorly documented for species with complex life-histories due to the difficulty of tracking dispersing individuals. Here we investigate variability in dispersal histories of a freshwater fish, Awaous stamineus, across the species' entire geographic range in the Hawaiian archipelago. Like many animals endemic to tropical island streams, these gobies have an amphidromous life cycle in which a brief marine larval phase enables dispersal among isolated freshwater habitats. Using otolith microchemistry, we document three distinct marine dispersal pathways, all of which are observed on every island. Surprisingly, we also find that 62% of individuals complete their life cycle entirely within freshwater, in contrast to the assumption that amphidromy is obligate in Hawaiian stream gobies. Comparing early life history outcomes based on daily otolith growth rings, we find that individuals with marine dispersal have shorter larval durations and faster larval growth, and their growth advantage over purely-freshwater counterparts continues to some degree into adult life. These individual benefits of maintaining a marine dispersal phase presumably balance against the challenge of finding and re-entering an island stream from the ocean. The facultative nature of amphidromy in this species highlights the selective balance between costs and benefits of dispersal in life history evolution. Accounting for alternative dispersal strategies will be essential for conservation of the amphidromous species that often dominate tropical island streams, many of which are at risk of extinction.
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  • 48
    Publication Date: 2014-03-13
    Description: Ecology, Volume 0, Issue 0, Ahead of Print. Climate change forecasts of more frequent climate extremes suggest that such events will become increasingly important drivers of future ecosystem dynamics and function. Because the rarity and unpredictability of naturally occurring climate extremes limits assessment of their ecological impacts, we experimentally imposed extreme drought and a mid-summer heat wave over two years in a central US grassland. While the ecosystem was resistant to heat waves, it was not resistant to extreme drought, which reduced aboveground net primary productivity (ANPP) below the lowest level measured in this grassland in almost thirty years. This extreme reduction in ecosystem function was a consequence of reduced productivity of both C4 grasses and C3 forbs. However, the dominant forb was negatively impacted by the drought more than the dominant grass, and this led to a reordering of species abundances within the plant community. Although this change in community composition persisted post-drought, ANPP recovered completely the year after drought due to rapid demographic responses by the dominant grass, compensating for loss of the dominant forb. Overall, these results show that an extreme reduction in ecosystem function attributable to climate extremes (e.g., low resistance) does not preclude rapid ecosystem recovery. Given that dominance by a few species is characteristic of most ecosystems, knowledge of the traits of these species and their responses to climate extremes will be key for predicting future ecosystem dynamics and function.
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  • 49
    Publication Date: 2014-03-13
    Description: Ecological Applications, Volume 0, Issue 0, Ahead of Print. The American pika (Ochotona princeps) has become a species of concern for its sensitivity to warm temperatures and potential vulnerability to global warming. We explored the value of radiocarbon dating of fecal pellets to address questions of population persistence and timing of site extirpation. Carbon was extracted from pellets collected at 43 locations in the western Great Basin, USA, including 3 known occupied sites and 40 sites of uncertain status at range margins or where previous studies indicated the species is vulnerable. We resolved calibrated dates with high precision (within several years), most of which fell in the period of the mid-late 20th century "bomb curve." The two-sided nature of the bomb curve renders "far-" and "near-side" dates of equal probability, which are separated by 1-4 decades. We document methods for narrowing resolution to one age range, including stratigraphic analysis of vegetation collected from pika haypiles. No evidence was found for biases in atmospheric 14C levels due to fossil-derived or industrial CO2 contamination. Radiocarbon dating indicated that pellets can persist for 〉59 years; known-occupied sites resolved contemporary dates. Using combined evidence from field observations and radiocarbon-dating, and the Bodie Mountains (Mtns) as an example, we propose a historical biogeographic scenario for pikas in minor Great Basin mountain ranges adjacent to major cordillera wherein historical climate variability led to cycles of extirpation and re-colonization during alternating cool and warm centuries. Using this model to inform future dynamics for small ranges in biogeographic settings similar to the Bodie Mtns, CA, extirpation of pikas appears highly likely under directional warming trends projected for the next century, even while populations in extensive cordillera (e.g., Sierra Nevada, Rocky Mtns, Cascade Range) are likely to remain viable due to extensive, diverse, habitat and high connectivity.
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  • 50
    Publication Date: 2014-03-13
    Description: Ecology, Volume 0, Issue 0, Ahead of Print. The resource quality of the host has been shown to affect parasite transmission success, prevalence and virulence. Seasonal availability of environmental nutrients alters density and stoichiometric quality (carbon-nutrient ratios) of both producers and consumers, suggesting that nutrient availability may drive fluctuations in parasite prevalence patterns observed in nature. We examined the interactions between the population dynamics of a keystone herbivore Daphnia and its parasites, and their associations with water nutrient concentrations, resource quantity and quality and other environmental variables (temperature, pH, oxygen concentration) in a small lake, using general linear models. We found that the prevalence of two gut endoparasites were positively related to food source and quality as well as nitrogen content of Daphnia - whereas the prevalence of an epibiont and overall parasite species richness were negatively related to phosphorus content of Daphnia. When only endoparasite species richness was considered, no connections to nutrients were found. Daphnia density was not connected to parasites, but we found interactions between Daphnia fecundity and parasite prevalence. Overall, our results suggest that environmental nutrient concentrations and stoichiometric quality of the host have a potential to affect seasonality in parasite epidemics, but the connections between environmental carbon:nutrient ratios and parasite prevalence patterns are diverse and species-specific.
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  • 51
    Publication Date: 2014-03-13
    Description: Ecology, Volume 0, Issue 0, Ahead of Print. The Tibetan Plateau (TP) is experiencing high rates of climatic change. We present a novel combined mechanistic-bioclimatic modeling approach to determine (i) how changes in precipitation and temperature on the TP may impact net primary production (NPP) in four major biomes (forest, shrub, grass, desert); and (ii) if there exists a maximum rain use efficiency (RUEMAX) that represents a "boundary that constrain(s) site-level productivity and efficiency" (Huxman et al. 2004). We used a daily mechanistic ecosystem model to generate 40-y outputs using observed climatic data for scenarios of (i) decreased precipitation (25 to 100%); (ii) increased air temperature (1 to 6°C); (iii) simultaneous changes in both precipitation ( ± 50%, ± 25%) and air temperature (+1 to +6°C); and (iv) increased interannual variability (IAV) of precipitation (+1σ to +3σ, with fixed means). We fitted model output from these scenarios to Huxman et al.'s RUEMAX bioclimatic model, NPP = α +RUE•PPT (where α = intercept, RUE is rain use efficiency and PPT is annual precipitation). Based on these analyses we conclude that: (i) there is strong support (when not explicit, then trend-wise) for Huxman et al.'s assertion that biomes converge to a common RUEMAX during the driest years at a site, thus representing the boundary for highest rain use efficiency; (ii) the interactive effects of simultaneously decreasing precipitation and increasing temperature on NPP for the TP is smaller than might be expected from additive, single-factor changes in these drivers; and (iii) that increasing IAV of precipitation may ultimately have a larger impact on biomes of the Tibetan Plateau than changing amounts of rainfall and air temperature alone.
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  • 52
    Publication Date: 2014-03-13
    Description: Ecology, Volume 0, Issue 0, Ahead of Print. Niche differentiation is a major hypothesized determinant of species distributions, but its practical importance is heavily debated and underlying mechanisms are poorly understood. Trait-based approaches have been used to infer niche differentiation and predict species distributions. For understanding underlying mechanisms, individual traits should be scaled up to whole-plant performance, which hardly has been done. We measured seven key traits that are important for carbon and water balance for 37 tropical tree species. We used a process-based plant physiological model to simulate the carbon budget of saplings along gradients of light and water availability, and quantified the performance of the species in terms of their light compensation points (a proxy for shade tolerance), water compensation points (proxy for drought tolerance) and maximum carbon gain rates (proxy for potential growth rate). We linked species performances to their observed distributions (the realized niches) at two spatial scales in Bolivian lowland forests: along a canopy openness gradient at local scale (~1 km2) and along a rainfall gradient (1100-2200 mm/y) at regional (~1000 km) scale. We show that the water compensation point was the best predictor of species distributions along water and light resource gradients within and across tropical forests. A sensitivity analysis suggests that the stomatal regulation of minimum leaf water potentials, rather than stem hydraulic traits (sapwood area and specific conductivity), contributed to the species differences in the water compensation point of saplings. The light compensation point and maximum carbon gain - both driven by leaf area index and the leaf nitrogen concentration - also contributed to differential species distributions at the local scale, but not or only marginally at the regional scale. Trait-and-physiology-based simulations of whole-plant performance thus help to evaluate the possible roles of individual traits in physiological processes underlying species performance along environmental gradients. The development of such whole plant concepts will improve our ability to understand responses of plant communities to shifts in resource availability and stress under global change.
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  • 53
    Publication Date: 2014-03-13
    Description: Ecology, Volume 0, Issue 0, Ahead of Print. Birds and their population dynamics are often used to understand and document anthropogenic effects on biodiversity. Nest success is a critical component of the breeding output of birds in different environments; but to obtain the complete picture of how bird populations respond to perturbations, we also need an estimate of nest abundance or density. The problem is that raw counts generally underestimate actual nest numbers because detection is imperfect and because some nests may fail or fledge before being subjected to detection efforts. Here we develop a state-space superpopulation capture-recapture approach in which inference about detection probability is based on the age at first detection, as opposed to the sequence of re-detections in standard capture-recapture models. We apply the method to ducks in which 1) the age of the nests and their initiation dates can be determined upon detection and 2) the duration of the different stages of the breeding cycle is a priori known. We fit three model variants with or without assumptions about the phenology of nest initiation dates, and use simulations to evaluate the performance of the approach in challenging situations. In an application to blue-winged teal Anas discors breeding at study sites in North and South Dakota, USA, nesting stage (egg-laying or incubation) markedly influenced nest survival and detection probabilities. Two individual covariates (one binary covariate: presence of grazing cattle at the nest site, and one continuous covariate: Robel index of vegetation) had only weak effects. We estimated that 5-10% of the total number of nests were available for detection but missed by field crews. An additional 6-15% were never available for detection. These proportions are expected to be larger in less intense, more typical sampling designs. User-friendly software nestAbund is provided to assist users in implementing the method.
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  • 54
    Publication Date: 2014-03-14
    Description: Ecology, Volume 0, Issue 0, Ahead of Print. Disease is often expected to limit host populations, but diseases do not always dramatically reduce host numbers and often have no effect. The impact of a fungal pathogen (Entomophaga grylli pathotype 1) on grasshopper (Camnula pellucida) populations was studied in a field experiment. We tested whether the effects of disease on grasshopper survival were 1) additive, with disease mortality summing with other mortality sources to determine the total population mortality rate or 2) compensatory, where disease mortality simply replaces mortality from other sources so that the total population mortality rate remains unchanged. We examined grasshopper survival in relation to differences in disease exposure, host density levels, and host developmental stage. The effects of disease varied with grasshopper developmental stage and density. Disease mortality increased by 60% at high grasshopper density compared to low density treatments, and decreased when grasshoppers fully matured. Despite increased rates of disease mortality at high densities, the total mortality rate was not notably higher in diseased grasshoppers (87%) compared to disease-free counterparts at high densities (83%), indicating that a large proportion of disease mortality simply replaced mortality from food limitation. Additive responses were supported in early and late instars, with disease exposure resulting in decreased grasshopper survival. In contrast, the effect of disease on adults was inconclusive. Yet, the disease did not affect adult survival suggesting that adult disease mortality is compensatory. Therefore, disease reduction of grasshopper populations (additive mortality) is more likely to occur during earlier developmental stages when hosts are most vulnerable to disease and at low host densities when food is abundant. Combined, our results emphasize the importance of host dynamics and food availability in how this host-pathogen system responds to disease. Accordingly, compensatory versus additive mortality may need to be considered when examining how disease ultimately affects host population dynamics.
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  • 55
    Publication Date: 2014-03-14
    Description: Ecology, Volume 0, Issue 0, Ahead of Print. Urbanization and global climate change can profoundly alter biological systems, yet scientists often analyze their effects separately. We test how the timing of life cycle events (phenology) is jointly influenced by these two components of global change. To do so, we use a long-term phenological dataset of 20 common butterfly species from 83 sites across the state of Ohio, USA, with sites that range from rural undeveloped areas to moderately sized cities. These sites span a several °C latitudinal gradient in mean temperature, mimicking the range of projected global climate warming effects through the end of the century. Although shifts toward earlier phenology are typical of species' responses to either global climate change or urbanization, we found that their interaction delayed several Ohio butterfly species' first appearance and peak abundance phenology. Exploitative species exhibited smaller delays in first appearance and peak abundance phenology in areas that were urbanized and geographically warm. Our results show that phenological responses to urbanization are contingent upon geographic variation in temperature, and that the impacts of urbanization and global climate change should be considered simultaneously when developing forecasts of biological responses to environmental change.
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  • 56
    Publication Date: 2014-03-14
    Description: Ecology, Volume 0, Issue 0, Ahead of Print. Fertilization via agricultural inputs and nutrient deposition is one of the major threats to global terrestrial plant richness, yet we still do not fully understand the mechanisms by which fertilization decreases plant richness. Tall clonal species have recently been proposed to cause declines in plant species richness by increasing in abundance in response to fertilization and competing strongly with other species. We tested this hypothesis in a fertilization experiment in a low productivity grassland by using a novel experimental manipulation of the presence vs. absence of clonal species and by examining the role of height within these treatments. We found that fertilization decreased species richness more in the presence than absence of clonal species. We also found that only tall species increased in biomass in response to fertilization. In the absence of clonal species, fertilization increased biomass of tall non-clonal species. However, in the presence of clonal species, fertilization decreased tall non-clonal biomass and only tall clonal biomass increased. Fertilization caused almost all short species to be lost in the presence, but not the absence, of clonal species and caused greater declines in the mean and variance of light levels in the presence of clonal species. These results show that the traits of species in a community can determine the magnitude of species loss due to fertilization. The strongly negative effect of tall clonals on species richness in fertilized plots is likely a result of their capacity to decrease light levels to a greater extent and more uniformly than non-clonal species, and thereby drive the exclusion of short species. These results help clarify the mechanisms whereby fertilization decreases grassland plant species richness and suggest that efforts to prevent the loss of species under fertilized conditions may be most effective when they focus on controlling the biomass of tall clonal species.
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  • 57
    Publication Date: 2014-03-14
    Description: Ecological Applications, Volume 0, Issue 0, Ahead of Print. Studies of predator-prey demographic responses and the physical drivers of such relationships are rare, yet essential for predicting future changes in the structure and dynamics of marine ecosystems. Here, we hypothesize that predator-prey relationships vary spatially in association with underlying physical ocean conditions, leading to observable changes in demographic rates, such as reproduction. To test this hypothesis, we quantified spatio-temporal variability in hydrographic conditions, krill, and forage fish to model predator (seabird) demographic responses over 18 years (1990-2007). We used principal component analysis and spatial correlation maps to assess coherence among ocean conditions, krill, and forage fish, and generalized additive models to quantify interannual variability in seabird breeding success relative to prey abundance. The first principal component of 4 hydrographic measurements yielded an index that partitioned "weak/warm upwelling" and "strong/cool upwelling" years. Partitioning of krill and forage fish time series among shelf and oceanic regions yielded spatially-explicit indicators of prey availability. Krill abundance within the oceanic region was remarkably consistent between years, whereas krill over the shelf showed marked interannual fluctuations in relation to ocean conditions. Anchovy abundance varied on the shelf, and was greater in years of strong stratification/weak upwelling and warmer temperatures. Spatio-temporal variability of juvenile forage fish covaried strongly with each other and with krill, but was weakly correlated with hydrographic conditions. Demographic responses between seabirds and prey availability revealed spatially-variable associations indicative of the dynamic nature of habitat-prey relationships. Quantification of spatially-explicit demographic responses, and their variability through time, demonstrates the possibility of delineating specific critical areas where the implementation of protective measures could maintain functions and productivity of central place foraging predators.
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  • 58
    Publication Date: 2014-03-14
    Description: We report a novel oscillatory pressure-assisted hot-pressing process for preparing high-quality ceramics. Compared with the samples prepared by conventional pressureless sintering (PS) and hot-pressing (HP), the zirconia ceramic prepared by oscillatory pressure-assisted hot-pressing (OPAHP) exhibited a higher density, smaller grain size, and more homogeneous structure. More remarkably, the strength of the OPAHP sample reached 1556 MPa, which is much higher than the samples prepared by other two techniques. The results suggest that OPAHP is a more effective technique for preparing high-quality zirconia, which is likely applicable to other material systems.
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  • 59
    Publication Date: 2014-05-02
    Description: The formation of a homogeneous Bi 8 TiO 14 phase was successfully achieved in a specimen calcined at 600°C. However, a Bi 4 Ti 3 O 12 secondary phase also developed in specimens calcined at temperatures higher than 600°C, probably because of Bi 2 O 3 evaporation. For specimens sintered above 800°C, a small amount of the Bi 8 TiO 14 phase melted during sintering, with the liquid phase contributing to the densification of the specimens; however, Bi 4 Ti 3 O 12 and Bi 12 TiO 20 secondary phases were still formed in these specimens. The microwave dielectric properties of the Bi 8 TiO 14 phase were considerably affected by variations in the microstructure of the specimens. When the sintering temperature exceeded 825°C, the amount of secondary phases increased, and this decreased the density and Q×f values of the specimens. Bi 8 TiO 14 ceramics sintered at 825°C exhibited promising microwave dielectric properties, with ε r = 47.4, Q×f  =   5370 GHz, and τ f = −16.01 ppm/°C.
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  • 60
    Publication Date: 2014-05-06
    Description: Ecological Applications, Ahead of Print. Translocation experiments, in which researchers displace animals then monitor their movements to return home, are commonly used as tools to assess functional connectivity of fragmented landscapes. Such experiments are purported to have important advantages of being time efficient and standardizing 'motivation' to move across individuals. Yet, we lack tests of whether movement behavior of translocated birds reflects natural behavior of unmanipulated birds. We compared the routine movement behavior of a tropical hummingbird (Phaethornis guy) to that of experimentally translocated individuals. We tested for differences in site-selection patterns during movement at two spatial scales (point and path levels). We also compared movement rates between treatments. Behaviors documented during translocation experiments reflected those observed during routine movements. At the point level, both translocated and non-translocated birds showed similar levels of preference for mature tropical forest. At the path level, step selection functions showed both translocated and non-translocated hummingbirds avoiding movement across non-forested matrix and selecting streams as movement corridors. Movement rates were generally higher during translocation experiments. However, the negative influence of forest cover on movement rates was proportionately similar in translocation and routine movement treatments. We report the first evidence showing that movement behavior of birds during translocation experiments is similar to their natural movement behavior. Therefore, translocation experiments may be reliable tools to address effects of landscape structure on animal movement. We observed consistent selection of landscape elements between translocated and non-translocated birds, indicating that both routine and translocation movement studies lead to similar conclusions regarding the effect of landscape structure and forest composition on functional connectivity. Our observations that hummingbirds avoid non-forest matrix and select riparian corridors also provides a potential mechanism for pollen limitation in fragmented tropical forest.
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  • 61
    Publication Date: 2014-05-06
    Description: Ecological Applications, Ahead of Print. Climate change vulnerability assessments for species of conservation concern often use species distribution and ecological niche modeling to project changes in habitat. One of many assumptions of these approaches is that food web dependencies are consistent in time and environmental space. Species at higher trophic levels that rely on the availability of species at lower trophic levels as food may be sensitive to extinction cascades initiated by changes in the habitat of key food resources. Here we assess climate change vulnerability for Ursus arctos (grizzly bears) in the southern Canadian Rocky Mountains using projected changes to 17 of the most commonly consumed plant food items. We used presence-absence information from 7,088 field plots to estimate ecological niches and to project changes in future distributions of each species. Model projections indicated idiosyncratic responses among food items. Many food items persisted or even increased, although several species were found to be vulnerable based on declines or geographic shifts in suitable habitat. This included Hedysarum alpinum (alpine sweet vetch), a critical spring and autumn root-digging resource when little else is available. Potential habitat loss was also identified for three fruiting species of lower importance to bears: Empetrum nigrum (crowberry), Vaccinium scoparium (grouseberry) and Fragaria virginiana (strawberry). A general trend towards uphill migration of bear foods may result in higher vulnerability to bear populations at low elevations which are also those that are most likely to have human-bear conflict problems. Regardless, a wide diet breadth of grizzly bears, as well as wide environmental niches of most food items, make climate change a much lower threat to grizzly bears than other bear species such as polar bears and panda bears. We cannot exclude, however, future alterations in human behavior and land use resulting from climate change that may reduce survival rates.
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  • 62
    Publication Date: 2014-05-06
    Description: Ecological Applications, Ahead of Print. Efforts to test and improve terrestrial biosphere models (TBMs) using a variety of data sources have become increasingly common. However, geographically extensive forest inventories have been under-exploited in previous model-data fusion efforts. Inventory observations of forest growth, mortality, and biomass integrate processes across a range of time scales, including slow time-scale processes such as species turnover, that are likely to have important effects on ecosystem responses to environmental variation. However, the large number (thousands) of inventory plots precludes detailed measurements at each location, so that uncertainty in climate, soil properties, and other environmental drivers may be large. Errors in driver variables, if ignored, introduce bias into model-data fusion. We estimated errors in climate and soil drivers at U.S. Forest Inventory and Analysis (FIA) plots, and we explored the effects of these errors on model-data fusion with the Geophysical Fluid Dynamics Laboratory LM3V dynamic global vegetation model. When driver errors were ignored or assumed small at FIA plots, responses of biomass production in LM3V to precipitation and soil available water capacity appeared steeper than the corresponding responses estimated from FIA data. These differences became non-significant if driver errors at FIA plots were assumed large. Ignoring driver errors when optimizing LM3V parameter values yielded estimates for fine-root allocation that were larger than biometric estimates, which is consistent with the expected direction of bias. To explore if complications posed by driver errors could be circumvented by relying on intensive study sites where driver errors are small, we performed a power analysis. To accurately quantify the response of biomass production to spatial variation in mean annual precipitation within the eastern U.S. would require at least 40 intensive study sites, which is larger than the number of sites typically available for individual biomes in existing plot networks. Driver errors may be accommodated by several existing model-data fusion approaches, including hierarchical Bayesian methods and ensemble filtering methods; however, these methods are computationally expensive. We propose a new approach, in which the TBM functional response is fit directly to the driver-error-corrected functional response estimated from data, rather than to the raw observations.
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  • 63
    Publication Date: 2014-05-06
    Description: Ecological Applications, Ahead of Print. Despite intensive monitoring, temporary emigration from the sampling area can induce bias severe enough for managers to discard life-history parameter estimates toward the terminus of the times series (terminal bias). Under random temporary emigration unbiased parameters can be estimated with CJS models. However, unmodeled Markovian temporary emigration causes bias in parameter estimates and an unobservable state is required to model this type of emigration. The robust design is most flexible when modeling temporary emigration, and partial solutions to mitigate bias have been identified, nonetheless there are conditions were terminal bias prevails. Long-lived species with high adult survival and highly variable non-random temporary emigration present terminal bias in survival estimates, despite being modeled with the robust design and suggested constraints. Because this bias is due to uncertainty about the fate of individuals that are undetected toward the end of the time series, solutions should involve using additional information on survival status or location of these individuals at that time. Using simulation, we evaluated the performance of models that jointly analyze robust design data and an additional source of ancillary data (predictive covariate on temporary emigration, telemetry, dead recovery, or auxiliary resightings) in reducing terminal bias in survival estimates. The auxiliary resighting and predictive covariate models reduced terminal bias the most. Additional telemetry data was effective at reducing terminal bias only when individuals were tracked for a minimum of two years. High adult survival of long-lived species made the joint model with recovery data ineffective at reducing terminal bias because of small-sample bias. The naïve constraint model (last and penultimate temporary emigration parameters made equal), was the least efficient, though still able to reduce terminal bias when compared to an unconstrained model. Joint analysis of several sources of data improved parameter estimates and reduced terminal bias. Efforts to incorporate or acquire such data should be considered by researchers and wildlife managers, especially in the years leading up to status assessments of species of interest. Simulation modeling is a very cost effective method to explore the potential impacts of using different sources of data to produce high quality demographic data to inform management.
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  • 64
    Publication Date: 2014-05-06
    Description: Ecological Applications, Ahead of Print. Warmer and drier climate over the past few decades has brought larger fire sizes and increased annual area burned in forested ecosystems of western North America, and continued increases in annual area burned are expected due to climate change. As warming continues, fires may also increase in severity and produce larger contiguous patches of high severity. We use remotely sensed burn-severity data from 125 fires in the northern Cascade Range of Washington, USA, to explore relationships between fire size, severity, and the spatial pattern of severity. We examine relationships between climate and the annual area burned and the size of wildfires over a 25-year period. We test the hypothesis that increased fire size is commensurate with increased burn severity and increased spatial aggregation of high severity. We also ask how local ecological controls might modulate these relationships by comparing results over the whole study area (the northern Cascade Range) to those from four ecological subsections within it. We found significant positive relationships between climate and fire size, and between fire size and the proportion of high severity and spatial-pattern metrics that quantify the spatial aggregation of high-severity areas within fires, but the strength and significance of these relationships varied among the four subsections. In areas with more contiguous subalpine forests and less complex topography, the proportion of high severity and spatial aggregation of high severity were more strongly correlated with fire size. If fire sizes increase in a warming climate, changes in the extent, severity, and spatial pattern of fire regimes will likely be more pronounced in higher-severity fire regimes with less complex topography and more continuous fuels.
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  • 65
    Publication Date: 2014-05-06
    Description: Ecological Applications, Ahead of Print. Fremont cottonwood (Populus fremonti, Eckenwal.) is a foundation riparian tree species that drives community structure and ecosystem processes in southwestern U.S. ecosystems. Despite its ecological importance, little is known about the ecological and environmental processes that shape its genetic diversity, structure and landscape connectivity. Here, we combine molecular analyses of 82 populations including 1,312 individual trees dispersed over the species' geographical distribution. We reduced the dataset to 40 populations and 743 individuals to eliminate admixture with a sibling species, and used multi-variate restricted optimization and reciprocal causal modeling, to evaluate the effects of river network connectivity and climatic gradients on gene flow. Our results confirmed the following: First, gene flow of Fremont cottonwood is jointly controlled by the connectivity of the river network and gradients of seasonal precipitation. Second, gene flow is facilitated by mid-sized to large rivers, and is resisted by small streams and terrestrial uplands, with resistance to gene flow decreasing with river size. Third, genetic differentiation increases with cumulative differences in winter and spring precipitation. Our results, suggest that ongoing fragmentation of riparian habitats will lead to a loss of landscape level genetic connectivity, leading to increased inbreeding and the concomitant loss of genetic diversity in a foundation species. These genetic effects will cascade to a much larger community of organisms, some of which are threatened and endangered.
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  • 66
    Publication Date: 2014-05-06
    Description: Ecological Applications, Ahead of Print. Identifying determinants of the probability and intensity of infections is important for understanding the epidemiology of wildlife diseases, and for managing their impact on threatened species. Chytridiomycosis, caused by the fungal pathogen Batrachochytrium dendrobatidis, has decimated populations of some amphibians. However, recent studies have identified important environmental constraints on the disease, related to the pathogen's physiological tolerances. In this study, we identified several intrinsic and extrinsic determinants of the probability and intensity of chytrid infections for the threatened Growling Grass Frog (Litoria raniformis) in south-eastern Australia, and used mark-recapture to estimate the effect of chytrid infections on the probability of survival of these frogs. Water temperature and salinity had negative effects on both the probability and intensity of chytrid infections. We coupled models of the infection process with a model of the effect of chytrid infections on the probability of survival to assess variation in the impact of chytridiomycosis between wetlands with differing temperature and salinity profiles. Our results suggest that warm, saline wetlands may be refuges from chytridiomycosis for L. raniformis, and should be priorities for protection. Our results also suggest that management actions that increase water temperature (e.g. reducing canopy shading) and salinity (e.g. complementing inflows with groundwater) could be trialled to reduce the impacts of chytridiomycosis on this species. This and other recent studies highlight the value of research on environmental risk factors for chytridiomycosis.
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  • 67
    Publication Date: 2014-05-06
    Description: Ecological Applications, Ahead of Print. Terrestrial soil is a large reservoir of atmospherically deposited mercury (Hg). However, few studies have evaluated the accumulation of Hg in terrestrial ecosystems in the northeastern United States, a region which is sensitive to atmospheric Hg deposition. In this study, we characterize Hg and organic matter in soil profiles from 139 sampling sites for five sub-regions across the northeastern United States, and estimate atmospheric Hg deposition to these sites by combining numerical modeling with experimental data from the literature. We did not observe any significant relationships between current net atmospheric Hg deposition and soil Hg concentrations or pools, even though soils are a net sink for Hg inputs. Soil Hg appears to be preserved relative to organic carbon (OC) and/or nitrogen (N) in the soil matrix, as a significant negative relationship was observed between the ratios of Hg/OC and OC/N (r = 0.54, p 〈 0.0001) that shapes the horizonal distribution patterns. We estimated that atmospheric Hg deposition since 1850 (3.97 mg m-2) accounts for 102% of the Hg pool in the organic horizons (3.88 mg m-2) and 19% of the total soil Hg pool (21.32 mg m-2), except for the Southern New England (SNE) sub-region. The mean residence time for soil Hg was estimated to be 1,800 years, except SNE which was 800 years. These patterns suggest that in additional to atmospheric deposition, the accumulation of soil Hg is linked to the mineral diagenetic and soil development processes in the region.
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  • 68
    Publication Date: 2014-05-06
    Description: Ecology, Ahead of Print. Predicting the future trend and viability of populations is an essential task in ecology. Because many populations respond to changing environments, uncertainty surrounding environmental responses must be incorporated into population assessments. However, understanding the effects of environmental variation on population dynamics requires information on several important demographic parameters which are often difficult to estimate. Integrated population models facilitate the integration of time-series data on population size and all existing demographic information from a species, allowing the estimation of demographic parameters for which limited or no empirical data exist. Although these models are ideal for assessments of population viability, they have so far not included environmental uncertainty. We incorporated environmental variation in an integrated population model to account for both demographic and environmental uncertainty in an assessment of population viability. In addition, we used this model to estimate true juvenile survival, an important demographic parameter for population dynamics that is difficult to estimate empirically. We applied this model to assess the past and future population trend of a rare island endemic songbird, the Montserrat Oriole Icterus oberi, which is threatened by volcanic activity. Montserrat Orioles experienced lower survival in years with volcanic ash-fall, causing periodic population declines that were compensated by higher seasonal fecundity in years with high pre-breeding season rainfall. Due to the inclusion of both demographic and environmental uncertainty in the model, the estimated population growth rate in the immediate future was highly imprecise (95% credible interval 0.844 - 1.105), and the probability of extinction after three generations (in the year 2028) was low (2.1%). This projection demonstrates that accounting for both demographic and environmental sources of uncertainty provides a more realistic assessment of the viability of populations under unknown future environmental conditions.
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  • 69
    Publication Date: 2014-05-06
    Description: Ecology, Ahead of Print. Grasslands have been lost and degraded in the USA since Euro-American settlement due to agriculture, development, introduced invasive species, and changes in fire regimes. Fire is frequently used in prairie restoration to control invasion by trees and shrubs, but may have additional consequences. For example, fire might reduce damage by herbivore and pathogen enemies by eliminating litter, which harbors eggs and spores. Less obviously, fire might influence enemy loads differently for native and introduced plant hosts. We used a controlled burn in a Willamette Valley (Oregon) prairie to examine these questions. We expected that without fire, introduced host plants should have less damage than native host plants because the introduced species are likely to have left many of their enemies behind when they were transported to their new range (the enemy release hypothesis, or ERH). If the ERH holds, then fire, which should temporarily reduce enemies on all species, should give an advantage to the natives because they should see greater total reduction in damage by enemies. Prior to the burn, we censused herbivore and pathogen attack on eight plant species (five of non-native origin: Bromus hordaceous, Cynosuros echinatus, Galium divaricatum, Schedonorus arundinaceus (=Festuca arundinacea), Sherardia arvensis; and three natives: Danthonia californica, Epilobium minutum, and Lomatium nudicale). The same plots were monitored for two years post-fire. Prior to the burn, native plants had more kinds of damage and more pathogen damage than introduced plants, consistent with the ERH. Fire reduced pathogen damage relative to the controls more for the native than the introduced species, but the effects on herbivory were negligible. Pathogen attack was correlated with plant reproductive fitness, whereas herbivory was not. These results suggest that fire may be useful for promoting some native plants in prairies due to its negative effects on their pathogens.
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  • 70
    Publication Date: 2014-05-06
    Description: Ecology, Ahead of Print. Floral nectar of many plant species is prone to colonization by microbial organisms such as yeasts. Their presence and metabolism of nectar chemical components have the potential to modify a suite of floral traits important for pollinator attraction, including nectar quality and scent. However, studies on the direct and indirect effects of nectar-inhabiting microogranisms on pollinator behavior and plant reproductive success remain rare. To determine their potential to affect pollinator behavior and plant fitness, we experimentally manipulated the common nectar-inhabiting yeast Metschnikowia reukaufii in the nectar of Delphinium nuttallianum, a short-lived montane perennial herb. We detected positive indirect, pollinator-mediated effects of yeasts on male plant fitness measured as pollen donation using powdered fluorescent dyes. However, we detected no direct or indirect effects on components of female fitness. Matching effects on male plant fitness, pollinators responded positively to the presence of yeasts, removing more nectar from flowers treated with M. reukaufii. Our results provide evidence of effects of nectar-inhabiting yeasts on male plant fitness and highlight the importance of microorganisms in mediating plant-pollinator interactions and subsequent plant fitness.
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  • 71
    Publication Date: 2014-05-06
    Description: Ecology, Ahead of Print. Complementary resource use and redundancy of species that fulfill the same ecological role are two mechanisms that can respectively increase and stabilize process rates in ecosystems. For example, predator complementarity and redundancy can determine prey consumption rates and their stability, yet few studies take into account the multiple predator species attacking multiple prey at different rates in natural communities. Thus, it remains unclear whether these biodiversity mechanisms are important determinants of consumption in entire predator-prey assemblages, such that food-web interaction structure determines community-wide consumption and stability. Here, we use empirical quantitative food-webs to study the community-wide effects of functional complementarity and redundancy of consumers (parasitoids) on herbivore control in temperate forests. We find that complementarity in host resource use by parasitoids was a strong predictor of absolute parasitism rates at the community level, and that redundancy in host-use patterns stabilised community-wide parasitism rates in space, but not through time. These effects can potentially explain previous contradictory results from predator diversity research. Phylogenetic diversity (measured using taxonomic distance) did not explain functional complementarity or parasitism rates, so could not serve as a surrogate measure for functional complementarity. Our study shows that known mechanisms underpinning predator diversity effects on both functioning and stability can easily be extended to link food webs to ecosystem functioning.
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  • 72
    Publication Date: 2014-05-06
    Description: Ecology, Ahead of Print. Understanding the way in which species are associated in communities is a fundamental question in ecology. Yet there remains a tension between communities as highly structured units or as coincidental collections of individualistic species. We explore these ideas using a new statistical approach that clusters species based on their environmental response- a species archetype, rather than clustering sites based on their species composition. We find that there are groups of species, which are consistently highly correlated, but that these groups are not unique to any set of locations and overlap spatially. The species present at a single site are a realisation of species from the (multiple) archetype groups that are likely to be present at that location based on their response to the environment.
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  • 73
    Publication Date: 2014-05-06
    Description: Ecology, Ahead of Print. Invasive alien predators can impose strong selection on native prey populations and induce rapid evolutionary change in the invaded communities. However, studies on evolutionary responses to invasive predators are often complicated by the lack of replicate populations differing in coexistence time with the predator, which would allow determining how prey traits change during the invasion. The red swamp crayfish Procambarus clarkii has invaded many freshwater areas worldwide, with negative impacts for native fauna. Here, we examined how coexistence time shapes antipredator responses of the Iberian waterfrog (Pelophylax perezi) to the invasive crayfish by raising tadpoles from five populations differing in historical exposure to P. clarkii (30 years, 20 years or no coexistence). Tadpoles from non-invaded populations responded to the presence of P. clarkii with behavioral plasticity (reduced activity), whereas long-term invaded populations showed canalized antipredator behavior (constant low activity level). Tadpoles from one of the long-term invaded populations responded to the crayfish with inducible morphological defenses (deeper tails), reflecting the use of both constitutive and inducible antipredator defenses against the exotic predator by this population. Our results suggest that, while naive P. perezi populations responded behaviorally to P. clarkii, the strong predation pressure imposed by the crayfish has induced the evolution of qualitatively different antipredator defenses in populations with longer coexistence time. These responses suggest that strong selection by invasive predators may drive rapid evolutionary change in invaded communities. Examining responses of prey species to biological invasions using multiple populations will help us better forecast the impact of invasive predators in natural communities.
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  • 74
    Publication Date: 2014-05-06
    Description: Ecology, Ahead of Print. Imbalances in phosphorus (P) intake relative to demand negatively affect animal growth, but their consequences are less understood for vertebrates, in which bone represents a significant and potentially flexible pool of P. Flexibility in body-P content could buffer vertebrates from the effects of imbalances between P intake and demand, reducing the likelihood of a sharp stoichiometric "knife-edge" in the relationship between growth rate and diet-P level. We conducted a meta-analysis of published aquaculture experiments that tested effects of diet %P on fish growth rate (49 studies, 28 species) and body-P content (27 of the studies in the main data set, 20 species). Our meta-analysis revealed significant P limitation of growth, as well as significant negative effects of excess P on growth rate. Diet-P thresholds for these effects occurred at ecologically relevant levels (mean ± SD optimal diet-P of 1.2 ± 0.45% under experimental conditions of high ration). Finally, the analysis also suggested a pattern of relatively shallow relationships between growth rate and diet-P level, coupled with surprisingly flexible body-P content in fishes. This result is consistent with fish using flexible body-P content (presumably mediated through bone P) to buffer imbalances between P intake and demand. Together, our results provide evidence for a relatively "dull" stoichiometric "knife-edge" in fishes, driven in part by flexible body-P content.
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  • 75
    Publication Date: 2014-05-06
    Description: Ecological Applications, Ahead of Print. Predicting how climate change is likely to interact with myriad other stressors that threaten species of conservation concern is an essential challenge in aquatic ecosystems. This study provides a framework to accomplish this task in salmon-bearing streams of the northwestern United States, where land-use related reductions in riparian shading have caused changes in stream thermal regimes, and additional warming from projected climate change may result in significant losses of coldwater fish habitat over the next century. Predatory non-native smallmouth bass have also been introduced into many northwestern streams and their range is likely to expand as streams warm, presenting an additional challenge to the persistence of threatened Pacific salmon. The goal of this work was to forecast the interactive effects of climate change, riparian management, and non-native species on stream-rearing salmon, and to evaluate the capacity of restoration to mitigate these effects. We intersected downscaled global climate forecasts with a local-scale water temperature model to predict mid- and end-of-century temperatures in streams in the Columbia River basin; we compared one stream that is thermally impaired due to the loss of riparian vegetation and another that is cooler and has a largely intact riparian corridor. Using the forecasted stream temperatures in conjunction with fish-habitat models, we predicted how stream-rearing Chinook salmon and bass distributions would change as each stream warmed. In the highly modified stream, end-of-century warming may cause near total loss of Chinook salmon rearing habitat and a complete invasion of the upper watershed by bass. In the less modified stream, bass were thermally restricted from the upstream-most areas. In both systems, temperature increases resulted in higher predicted spatial overlap between stream-rearing Chinook salmon and potentially predatory bass in the early summer (2-4-fold increase) and greater abundance of bass. We found that riparian restoration could prevent the extirpation of Chinook salmon from the more altered stream, and could also restrict bass from occupying the upper 31 km of salmon rearing habitat. The proposed methodology and model predictions are critical for prioritizing climate-change adaptation strategies before salmonids are exposed to both warmer water and greater predation risk by non-native species.
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  • 76
    Publication Date: 2014-05-06
    Description: Ecological Applications, Volume 24, Issue 3, Page 528-538, April 2014. A unique high temporal frequency data set from an irrigated cotton–wheat rotation was used to test the agroecosystem model DayCent to simulate daily N2O emissions from subtropical vertisols under different irrigation intensities. DayCent was able to simulate the effect of different irrigation intensities on N2O fluxes and yield, although it tended to overestimate seasonal fluxes during the cotton season. DayCent accurately predicted soil moisture dynamics and the timing and magnitude of high fluxes associated with fertilizer additions and irrigation events. At the daily scale we found a good correlation of predicted vs. measured N2O fluxes (r2 = 0.52), confirming that DayCent can be used to test agricultural practices for mitigating N2O emission from irrigated cropping systems. A 25-year scenario analysis indicated that N2O losses from irrigated cotton–wheat rotations on black vertisols in Australia can be substantially reduced by an optimized fertilizer and irrigation management system (i.e., frequent irrigation, avoidance of excessive fertilizer application), while sustaining maximum yield potentials.
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  • 77
    Publication Date: 2014-05-06
    Description: Ecological Applications, Ahead of Print. Grazing represents one of the most common disturbances in drylands worldwide, affecting both ecosystem structure and functioning. Despite the efforts to understand the nature and magnitude of grazing effects on ecosystem components and processes, contrasting results continue to arise. This is particularly remarkable for the biological soil crust (BSC) communities (i.e. cyanobacteria, lichens and bryophytes), which play an important role in soil dynamics. Here we evaluated simultaneously the effect of grazing impact on BSC communities (resistance) and recovery after livestock exclusion (resilience) in a semiarid grassland of Central Mexico. In particular, we examined BSC species distribution, species richness, taxonomical group cover (i.e., cyanobacteria, lichen, bryophyte) and composition along a disturbance gradient with different grazing regimes (low, medium, high impact) and along a recovery gradient with differently-aged livestock exclosures (short-, medium-, long-term exclusion). Differences in grazing impact and time of recovery from grazing both resulted in slight changes in species richness; however, there were pronounced shifts in species composition and group cover. We found we could distinguish four highly diverse and dynamic BSC species groups: 1) species with high resistance and resilience to grazing, 2) species with high resistance but low resilience, 3) species with low resistance but high resilience, and 4) species with low resistance and resilience. While disturbance resulted in a novel diversity configuration, which may profoundly affect ecosystem functioning, we observed that 10 years of disturbance removal did not lead to the ecosystem structure found after 27 years of recovery. These findings are an important contribution to our understanding of BCS dynamics from a species and community perspective placed in a land use change context.
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  • 78
    Publication Date: 2014-05-06
    Description: Ecological Applications, Ahead of Print. Post-fire predictions of forest recovery under future climate change and management actions are necessary for forest managers to make decisions about treatments. We applied the Climate-Forest Vegetation Simulator (Climate-FVS), a new version of a widely used forest management model, to compare alternative climate and management scenarios in a severely burned multi-species forest of Arizona, U.S.A. The incorporation of seven combinations of General Circulation Models (GCM) and emissions scenarios altered long-term (100 years) predictions of future forest condition compared to a No Climate Change (NCC) scenario, which forecast a gradual increase to high levels of forest density and carbon stock. In contrast, emissions scenarios that included continued high greenhouse gas releases led to near-complete deforestation by 2111. GCM-emissions scenario combinations that were less severe reduced forest structure and carbon stock relative to NCC. Fuel reduction treatments that had been applied prior to the severe wildfire did have persistent effects, especially under NCC, but were overwhelmed by increasingly severe climate change. We tested six management strategies aimed at sustaining future forests: prescribed burning at 5, 10, or 20-year intervals, thinning 40% or 60% of stand basal area, and no-treatment. Severe climate change led to deforestation under all management regimes, but important differences emerged under the moderate scenarios: treatments that included regular prescribed burning fostered low density, wildfire-resistant forests composed of the naturally dominant species, ponderosa pine. Non-fire treatments under moderate climate change were forecast to become dense and susceptible to severe wildfire, with a shift to dominance by sprouting species. Current U.S. forest management requires modeling of future scenarios but does not mandate consideration of climate change effects. However, this study showed substantial differences in model outputs depending on climate and management actions. Managers should incorporate climate change into the process of analyzing the environmental effects of alternative actions.
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  • 79
    Publication Date: 2014-05-06
    Description: Ecological Applications, Ahead of Print. Digital repeat photography is becoming widely used for near surface remote sensing of vegetation. Canopy greenness, which has been used extensively for phenological applications, can be readily quantified from camera images. Important questions remain, however, as to whether the observed changes in canopy greenness are directly related to changes in leaf-level traits, changes in canopy structure, or some combination thereof. We investigated relationships between canopy greenness and various metrics of canopy structure and function, using five years (2008-2012) of automated digital imagery, ground observations of phenological transitions, leaf area index (LAI) measurements, and eddy-covariance estimates of gross ecosystem photosynthesis from the Harvard Forest, a temperate deciduous forest in the northeastern USA. Additionally, we sampled canopy sunlit leaves on a weekly basis throughout the growing season of 2011. We measured physiological and morphological traits including leaf size, mass (wet/dry), nitrogen content, chlorophyll fluorescence, and spectral reflectance, and characterized individual leaf color with flatbed scanner imagery. Our results show that observed spring and autumn phenological transition dates are well captured by information extracted from digital repeat photography. However, spring development of both LAI and the measured physiological and morphological traits are shown to lag behind spring increases in canopy greenness, which rises very quickly to its maximum value before leaves are even half their final size. Based on the hypothesis that changes in canopy greenness represent the aggregate effect of changes in both leaf-level properties (specifically, leaf color) and changes in canopy structure (specifically, LAI), we developed a two end-member mixing model. With just a single free parameter, the model was able to reproduce the observed seasonal trajectory of canopy greenness. This analysis shows that canopy greenness is relatively insensitive to changes in LAI at high LAI levels, which we further demonstrate by assessing the impact of an ice-storm on both LAI and canopy greenness. Our study provides new insights into the mechanisms driving seasonal changes in canopy greenness retrieved from digital camera imagery. The nonlinear relationship between canopy greenness and canopy LAI has important implications both for phenological research applications and for assessing responses of vegetation to disturbances.
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  • 80
    Publication Date: 2014-05-06
    Description: Ecology, Ahead of Print. Community ecologists use functional groups based on the rarely tested assumption that within-group responses to ecological processes are similar and thus members are functionally equivalent. However, recent research suggests functional equivalency may break down with human impacts. We tested the equivalency assumption and model predictions of responses to simulated human alterations in nutrients and large herbivores for two models of coral reef algae, the Relative Dominance Model (RDM) and the Functional Group Model (FGM). Results of both mesocosm and field experiments using assembled communities were compared to model predictions and within- and between-group variability were assessed. Both models' predictions of group response to herbivory matched experimental outcomes, but only the RDM predicted response to nutrients. However, within-group variability was dramatic, as the RDM grouped species with opposite responses to herbivory and the FGM grouped species with unique responses to nutrients. These heterogeneous responses resulted in loss of information and masked strong interactions between herbivory and nutrients that were not included in the models. As humans continue to impact major ecological processes in ecosystems globally, we postulate that functional-group models may need to be reformulated to account for shifting baselines.
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  • 81
    Publication Date: 2014-05-06
    Description: Ecology, Ahead of Print. Flowering plants often occur in mixed-species groups where interactions between them can occur both pre- and post-pollination. In post-pollination interactions conspecific (CP) and heterospecific pollen (HP) interact on the style and previous work indicates that these interactions, often, but not always, lead to reduced fitness. However, over half of insect-pollinated species display a mixed mating system, where both self and outcross CP could interact with HP, yet no study has evaluated the effects on both types of CP. We present and test hypotheses for the differential effects of HP depending on CP source and timing of HP arrival with a mixed-mating plant, M. guttatus, and one of its HP donors (Helianthus exilis). We found H. exilis pollen reduced tube growth and fertilization success of M. guttatus self pollen to a greater degree than outcross pollen, but only when CP and HP were applied simultaneously. Our results indicate that for mixed-mating plants the fitness consequences of HP receipt can be more detrimental than previously thought. Furthermore, our findings suggest that HP receipt, which can be high in biodiverse areas, has the potential to influence mating system and genetic diversity of the plant populations within them.
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  • 82
    Publication Date: 2014-05-06
    Description: Ecology, Ahead of Print. Factors that control tree seedling dynamics are critical determinants of forest diversity. We examined the role of density-dependent mortality and abiotic factors in the differential establishment and survival of tree seedlings at three large mapped forest plots in Indiana, Virginia and Wisconsin, USA. We tested whether seedling densities and seedling survival are related to local biotic and abiotic factors with generalized linear mixed models. Spatial point pattern analyses were utilized to determine if the distribution patterns of seedlings and saplings are consistent with a pattern generated by negative density-dependent mortality with respect to conspecific trees. Initial sampled seedling density for nearly a third of species showed a positive correlation with increasing conspecific basal area indicating dispersal limitation, but few had any association with abiotic variables. By contrast, survival of seedlings over one year significantly declined with increasing conspecific basal area. Point pattern analyses indicated that nearly one third of tree species had significantly over-dispersed point patterns of conspecific seedlings and saplings relative to adult densities; the majority of other species exhibited random spatial arrangements. Our results demonstrate that negative conspecific density-dependent mortality of seedlings could generate the spatial patterns observed at later life stages. By differentially favoring seedlings of other species, this process may contribute to the maintenance of tree diversity in temperate forests, just as others have demonstrated for tropical forests.
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  • 83
    Publication Date: 2014-05-06
    Description: Ecology, Ahead of Print. Communities are comprised of individual species that respond to changes in their environment depending in part on their niche requirements. These species comprise the biodiversity of any given community. Common biodiversity metrics such as richness, evenness, and the species abundance distribution are frequently used to describe biodiversity across ecosystems and taxonomic groups. While it is increasingly clear that researchers will need to forecast changes in biodiversity, ecology currently lacks a framework for understanding the natural background variability in biodiversity or how biodiversity patterns will respond to environmental change. We predict that while species populations depend on local ecological mechanisms (e.g., niche processes) and should respond strongly to disturbance, community-level properties that emerge from these species should generally be less sensitive to disturbance because they depend on regional mechanisms (e.g., compensatory dynamics). Using published data from terrestrial animal communities, we show that community-level properties were generally resilient under a suite of artificial and natural manipulations. In contrast, species responded readily to manipulation. Our results suggest that community-level measures are poor indicators of change, perhaps because many systems display strong compensatory dynamics maintaining community-level properties. We suggest that ecologists consider using multiple metrics that measure composition and structure in biodiversity response studies.
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  • 84
    Publication Date: 2014-05-06
    Description: Ecology, Ahead of Print. In colonial-breeding species, prebreeders often emigrate temporarily from natal reproductive colonies then subsequently return for one or more years before producing young. Variation in attendance/nonattendance patterns can have implications for subsequent recruitment. We used open robust-design multi-state models and 27 years of encounter data for prebreeding female Weddell seals (Leptonychotes weddellii Lesson) to evaluate hypotheses about (1) the relationships of temporary emigration (TE) probabilities to environmental and population size covariates, (2) motivations for attendance and consequences of nonattendance for subsequent probability of recruitment to the breeding population. TE probabilities were density-dependent (= 0.66, = 0.17) (βBPOP = 0.66, SE = 0.17)and increased when the fast-ice edge was distant from the breeding colonies (βDIST = 0.75, SE = 0.04), and were strongly age- and state-dependent. These results suggest that tradeoffs between potential benefits and costs of colony attendance vary annually and might influence motivation to attend colonies. Recruitment probabilities were greatest for seals that consistently attended colonies in 2 or more years (e.g.,ψage10 = 0.56, sd = 0.17) and lowest for seals that never or inconsistently attended prior to recruitment (e.g., ψage10 = 0.32, sd = 0.15). In colonial-breeding seabirds repeated colony attendance increases subsequent probability of recruitment to the adult breeding population; our results suggest similar implications for a marine mammal and are consistent with the hypothesis that prebreeders were motivated to attend reproductive colonies to gain reproductive skills or perhaps to optimally synchronize estrus through close association with mature breeding females.
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  • 85
    Publication Date: 2014-05-06
    Description: Ecology, Ahead of Print. Fertilization via agricultural inputs and nutrient deposition is one of the major threats to global terrestrial plant richness, yet we still do not fully understand the mechanisms by which fertilization decreases plant richness. Tall clonal species have recently been proposed to cause declines in plant species richness by increasing in abundance in response to fertilization and competing strongly with other species. We tested this hypothesis in a fertilization experiment in a low productivity grassland by using a novel experimental manipulation of the presence vs. absence of clonal species and by examining the role of height within these treatments. We found that fertilization decreased species richness more in the presence than absence of clonal species. We also found that only tall species increased in biomass in response to fertilization. In the absence of clonal species, fertilization increased biomass of tall non-clonal species. However, in the presence of clonal species, fertilization decreased tall non-clonal biomass and only tall clonal biomass increased. Fertilization caused almost all short species to be lost in the presence, but not the absence, of clonal species and caused greater declines in the mean and variance of light levels in the presence of clonal species. These results show that the traits of species in a community can determine the magnitude of species loss due to fertilization. The strongly negative effect of tall clonals on species richness in fertilized plots is likely a result of their capacity to decrease light levels to a greater extent and more uniformly than non-clonal species, and thereby drive the exclusion of short species. These results help clarify the mechanisms whereby fertilization decreases grassland plant species richness and suggest that efforts to prevent the loss of species under fertilized conditions may be most effective when they focus on controlling the biomass of tall clonal species.
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  • 86
    Publication Date: 2014-05-06
    Description: Ecology, Ahead of Print. Nutrient pollution constitutes a major threat to biodiversity - one whose magnitude is predicted to increase greatly in the near future. While the negative effects of excessive nitrogen and phosphorus loading on local species diversity are widely appreciated, a growing body of evidence indicates that increases in productivity (a correlate of nutrient supply) can also reduce predictability in community composition by driving community divergence and increases in beta diversity (or compositional dissimilarity among communities). Stochastic variation in dispersal history has frequently been cited as a cause of such patterns. However, underlying mechanisms have not received strong experimental scrutiny. I present results of a microcosm experiment testing the effects of enrichment and dispersal mode on zooplankton community structure. I show that beta diversity increases with enrichment but only in the presence of sequential dispersal and variation in dispersal history. Sequential dispersal and enrichment enhanced beta diversity by driving increases in temporal compositional turnover (or beta diversity in time). These results suggest that nutrient enrichment and dispersal may have interactive effects on community organization by facilitating persistent compositional flux and reducing our ability to predict the structure of communities in both space and time.
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  • 87
    Publication Date: 2014-05-06
    Description: Ecology, Ahead of Print. Climate change forecasts of more frequent climate extremes suggest that such events will become increasingly important drivers of future ecosystem dynamics and function. Because the rarity and unpredictability of naturally occurring climate extremes limits assessment of their ecological impacts, we experimentally imposed extreme drought and a mid-summer heat wave over two years in a central US grassland. While the ecosystem was resistant to heat waves, it was not resistant to extreme drought, which reduced aboveground net primary productivity (ANPP) below the lowest level measured in this grassland in almost thirty years. This extreme reduction in ecosystem function was a consequence of reduced productivity of both C4 grasses and C3 forbs. However, the dominant forb was negatively impacted by the drought more than the dominant grass, and this led to a reordering of species abundances within the plant community. Although this change in community composition persisted post-drought, ANPP recovered completely the year after drought due to rapid demographic responses by the dominant grass, compensating for loss of the dominant forb. Overall, these results show that an extreme reduction in ecosystem function attributable to climate extremes (e.g., low resistance) does not preclude rapid ecosystem recovery. Given that dominance by a few species is characteristic of most ecosystems, knowledge of the traits of these species and their responses to climate extremes will be key for predicting future ecosystem dynamics and function.
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  • 88
    Publication Date: 2014-05-06
    Description: Ecology, Ahead of Print. Thresholds profoundly affect our understanding and management of ecosystem dynamics, but we have yet to develop practical techniques to assess the risk that thresholds will be crossed. Combining ecological knowledge of critical system interdependencies with a large-scale experiment we tested for breaks in the ecosystem interaction network to identify threshold potential in real world ecosystem dynamics. Our experiment on marine sandflats demonstrated that reductions in incident sunlight changed the interaction network between sediment biogeochemical fluxes, productivity and macrofauna. By demonstrating loss of positive feedbacks and changes in the architecture of the network, we provide mechanistic evidence that stressors lead to breakpoints in dynamics, which theory predicts pre-dispose a system to a critical transition.
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  • 89
    Publication Date: 2014-05-06
    Description: Ecology, Ahead of Print. Pendulous lichens dominate canopies of boreal forests, with dark Bryoria species in upper versus light Alectoria and Usnea species in lower canopy. These genera offer important ecosystem services like winter forage for reindeer and caribou. The mechanism behind this niche separation is poorly understood. We tested the hypothesis that species-specific sun-screening fungal pigments protect underlying symbiotic algae differently against high light, and thus shape the vertical canopy gradient of epiphytes. Three pale species with the reflecting pigment usnic acid (Alectoria sarmentosa, Usnea dasypoga, U. longissima) and three with dark, absorbing melanins (Bryoria capillaris, B. fremontii, B. fuscescens) were compared. We subjected the lichens to desiccation stress with and without light, and assessed their performance with chlorophyll fluorescence. Desiccation alone only affected U. longissima. By contrast, light in combination with desiccation caused photoinhibitory damage in all species. Usnic lichens were significantly more susceptible to light during desiccation than melanic ones. Thus, melanin is a more efficient light-screening pigment than usnic acid. Thereby, the vertical gradient of pendulous lichens in forest canopies is consistent with a shift in type and functioning of sun-screening pigments, from high-light tolerant Bryoria in the upper to susceptible Alectoria and Usnea in the lower canopy.
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  • 90
    Publication Date: 2014-05-06
    Description: Ecology, Ahead of Print. Predicting the effects of climate change on ecological communities requires an understanding of how environmental factors influence both physiological processes and species interactions. Specifically, the net impact of temperature on community structure depends on the relative response of physiological energetic costs (metabolism) and energetic gains (ingestion of resources) that mediate the flow of energy throughout a food web. However, the relative temperature scaling of metabolic and ingestion rates have rarely been measured for multiple species within an ecological assemblage and it is not known how, and to what extent, these relative scaling differences vary among species. To investigate the relative influence of these processes, I measured the temperature scaling of metabolic and ingestion rates for a suite of rocky intertidal species using a multiple regression experimental design. I compared oxygen consumption rates (as a proxy for metabolic rate) and ingestion rates by estimating the temperature scaling parameter of the 'universal temperature dependence' (UTD) model, a theoretical model derived from first principles of biochemical kinetics and allometry. The results show that consumer metabolic rates (energetic costs) tend to be more sensitive to temperature than ingestion rates (energetic gains). Thus, as temperature increases, metabolic rates tend to increase faster relative to ingestion rates, causing the overall energetic efficiencies of these rocky intertidal invertebrates to decline. Metabolic and ingestion rates largely scaled in accordance with the UTD model; however, non-linearity was evident in several cases, particularly at higher temperatures, in which alternative models were more appropriate. There are few studies where multiple rate dependencies are measured on multiple species from the same ecological community. These results indicate that there may be general patterns across species in the temperature scaling of biological rates, with important implications for forecasting temperature effects on ecological communities.
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  • 91
    Publication Date: 2014-05-06
    Description: Ecology, Ahead of Print. Metacommunity theory generally predicts that regional dispersal of organisms among local habitat patches should influence spatial patterns of species diversity. In particular, increased dispersal rates are generally expected to increase local (α) diversity, yet homogenize local communities across the region (decreasing β-diversity), resulting in no change in regional (γ) diversity. Although the effect of dispersal on α-diversity has garnered much experimental attention, the influence of dispersal rates on diversity at larger spatial scales (β and γ) is poorly understood. Furthermore, these theoretical predictions are not well tested in the field, where other environmental factors (e.g., habitat size, resource density) likely also influence species diversity. Here, we use a system of freshwater rock pools on Appledore Island, Maine, USA to test the effects of dispersal rate on species diversity in metacommunities. The pools exist in clusters (metacommunities) that experience different levels of dispersal imposed by gulls (Larus spp.), which we show to be frequent passive dispersers of rock-pool invertebrates. Although previous research has suggested that waterbirds may disperse aquatic invertebrates, our study is the first to quantify the rate at which such dispersal occurs and determine its effects on species diversity. In accordance with theory, we found that metacommunities experiencing higher dispersal rates had significantly more homogeneous local communities (reduced β-diversity) and that γ-diversity was not influenced by dispersal rate. Contrary to theoretical predictions, however, α-diversity in the rock pools was not significantly influenced by dispersal. Rather, local diversity was significantly positively related to local habitat size and both α- and γ-diversity were influenced by the physicochemical environment of the pools. These results provide an important field test of metacommunity theory, highlighting how local and regional factors interact to drive patterns of species diversity in metacommunities, and demonstrate that waterbirds are indeed important dispersal vectors for aquatic invertebrates.
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  • 92
    Publication Date: 2014-05-06
    Description: Ecology, Ahead of Print. Given the health and economic burden associated with the widespread occurrence of co-infections in human and agricultural animals, understanding how co-infections contribute to host heterogeneity to infection and transmission is critical if we are to assess risk of infection based on host characteristics. Here, we examine whether host heterogeneity to infection leads to similar heterogeneity in transmission in a population of rabbits single and co-infected with two helminths and monitored monthly for eight years. Compared to single infections, co-infected rabbits carried higher Trichostrongylus retortaeformis intensities, longer worms with fewer eggs in utero and shed a similar numbers of parasite eggs. In contrast, the same co-infected rabbits harbored fewer Graphidium strigosum with longer bodies and more eggs in utero, and shed more eggs of this helminth. A positive density-dependent relationship between fecundity and intensity was found for T. retortaeformis but not G. strigosum in co-infected rabbits. Juvenile rabbits contributed to most of the infection and shedding of T. retortaeformis while adult hosts were more important for G. strigosum dynamics of infection and transmission, and this pattern was consistent in single and co-infected individuals. This host-parasite system suggests that we cannot predict pattern of parasite shedding during co-infections based on intensity of infection alone. We suggest that a mismatching between susceptibility and infectiousness should be expected in helminth co-infections and should not be overlooked.
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  • 93
    Publication Date: 2014-05-06
    Description: Ecology, Ahead of Print. Large pelagic predators play important roles in oceanic ecosystems and may migrate vast distances to utilize resources in different marine ecoregions. Understanding movement patterns of migratory marine animals is critical for effective management, but often challenging due to the cryptic habitat of pelagic migrators and the difficulty of assessing past movements. Chemical tracers can partially circumvent these challenges by reconstructing recent migration patterns. Pacific bluefin tuna (Thunnus orientalis; PBFT) inhabit the western and eastern Pacific Ocean and are in steep decline due to overfishing. Understanding age-specific eastward trans-Pacific migration patterns can improve management practices but these migratory dynamics remain largely unquantified. Here, we combine a Fukushima-derived radiotracer (134Cs) with bulk tissue- and amino acid-stable isotope analyses of PBFT to distinguish recent migrants from residents of the eastern Pacific Ocean. The proportion of recent migrants to residents decreased in older year classes, though the proportion of older PBFT that recently migrated across the Pacific was greater than previous estimates. This novel toolbox of biogeochemical tracers can be applied to any species that crosses the North Pacific Ocean.
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  • 94
    Publication Date: 2014-05-06
    Description: Ecology, Ahead of Print. The emission of volatile monoterpenes from coniferous trees impacts the oxidative state of the troposphere and multi-trophic signaling between plants and animals. Previous laboratory studies have revealed that climate anomalies and herbivory alter the rate of tree monoterpene emissions. However, no studies to date have been conducted to test these relations in situ. We conducted a two-year field experiment at two semi-arid sites dominated by pinyon pine (Pinus edulis) during outbreaks of a specialist herbivore, the southwestern tiger moth (Lophocampa ingens: Arctiidae). We discovered that during the early spring, when herbivory rates were highest, monoterpene emission rates were approximately two to six times higher from undamaged needles on damaged trees, with this increase in emissions due to α-pinene, β-pinene, and camphene at both sites. During mid-summer, emission rates did not differ between previously damaged and undamaged trees at the site on the Western Slope of the Rocky Mountains, but rather tracked changes in the temperature and precipitation regime characteristic of the region. As the mid-summer drought progressed at the Eastern Slope site, emission rates were low but differences between previously damaged and undamaged trees were not statistically significant. Despite no difference in emissions, mid-summer tissue monoterpene concentrations were significantly lower in previously damaged trees at both sites. With the onset of monsoon rains during late summer, emission rates from previously damaged trees increased to levels higher than those of undamaged trees despite the lack of herbivory. We conclude (1) herbivory systemically increases the flux of terpenes to the atmosphere during the spring, (2) drought overrides the effect of past herbivory as the primary control over emissions during the mid-summer, and (3) that a release from drought and the onset of late-summer rains is correlated with a secondary increase in emissions, particularly from herbivore-damaged trees, possibly due to a drought-delayed stimulation of induced monoterpene synthesis and/or increases in stomatal conductance. A greater understanding of the interactive effects of seasonality and herbivory on monoterpene emissions provides much needed information regarding the atmospheric and ecological consequences these compounds will have on semi-arid ecosystems.
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  • 95
    Publication Date: 2014-05-06
    Description: Ecology, Ahead of Print. Fires are the major natural disturbance in the boreal forest, and their frequency and intensity will likely increase as the climate warms. Terrestrial nutrients released by fires may be transported to boreal lakes, stimulating increased primary productivity, which may radiate through multiple trophic levels. Using a before-after-control-impact (BACI) design, with pre- and post-fire data from burned and unburned areas, we examined effects of a natural fire across several trophic levels of boreal lakes, from nutrient and chlorophyll levels to macroinvertebrates to waterbirds. Concentrations of total nitrogen and phosphorus were not affected by the fire. Chlorophyll a levels were also unaffected, likely reflecting the stable nutrient concentrations. For aquatic invertebrates, we found that densities of 3 functional feeding groups did not respond to the fire (filterers, gatherers, scrapers), while 2 groups increased (shredders, predators). Amphipods accounted for 98% of shredder numbers, and we hypothesize that fire-mediated habitat changes may have favored their generalist feeding and habitat ecology. This increase in amphipods may have in turn driven increased predator densities, as amphipods were the most numerous invertebrate in our lakes and are commonly taken as prey. Finally, abundance of waterbird young, which feed primarily on aquatic invertebrates, was not affected by the fire. Overall, ecosystems of our study lakes were largely resilient to forest fires, likely due to their high initial nutrient concentrations and small catchment sizes. Moreover, this resilience spanned multiple trophic levels, a significant result for ecologically similar boreal regions, especially given the high potential for increased fires with future climate change.
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  • 96
    Publication Date: 2014-05-06
    Description: Ecology, Ahead of Print. The importance of terrestrial-derived organic matter for lake zooplankton communities remains debated, partly because little is known about the basic pathways by which allochthonous carbon is transferred to zooplankton, and whether these vary among the major taxonomic and functional groups. We quantified allochthony of three zooplankton groups (Cladocera, Calanoida and Cyclopoida) across 18 lakes in Québec, spanning broad gradients of dissolved organic matter (DOM) and lake trophy, using a multi-isotope (δ2H + δ13C), multi-source (terrestrial, phytoplanktonic, benthic) approach. All three zooplankton groups had significant levels of allochthony, but differed greatly in their respective patterns across lakes. Allochthony in Calanoida and Cyclopoida was linked to detrital food chains based on particulate organic matter (POM) and on DOM, respectively, whereas in Cladocera it appeared related to both pathways; not surprisingly this latter group had the highest mean allochthony (0.31; compared to 0.18 in Cyclopoida and 0.16 in Calanoida). This study highlights the complexity of the pathways of delivery and transfer of terrestrial organic matter in freshwaters, and underscores the role that microbial food webs play in this transfer.
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  • 97
    Publication Date: 2014-05-06
    Description: Ecology, Ahead of Print. Consumer growth can be affected by imbalances between the nutrient content of the consumer and its food resource. Although ontogenetic-driven changes in animal composition are well-documented, their potential consequences for organism's sensitivity to food quality constraints have remained elusive. Here we show that the potential growth response of the copepod Mixodiaptomus laciniatus (as %RNA and RNA:DNA ratio) to the natural gradient of seston carbon (C):nutrient ratio is unimodal and stage-specific. Solution of the equation given by the first derivative function provided the optimum C:nutrient ratio for maximum stage-specific growth, which increased during ontogeny. The peakedness of the function indicated that animal vulnerability to suboptimal food quality decreased as juveniles reached adulthood. Consistent with these results, a field-experiment demonstrated that potential consumer growth responded to variations in seston C:phosphorus ratio and that early-life stages were particularly vulnerable to suboptimal food quality.
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  • 98
    Publication Date: 2014-05-06
    Description: Ecology, Ahead of Print. Soils host diverse communities of interacting microbes and the nature of interspecific interactions is increasingly recognised to affect ecosystem-level processes. Antagonistic interactions between bacteria and fungi are of particular relevance for soil functioning. A number of soil bacteria produce secondary metabolites that inhibit eukaryotic growth. Antibiosis may be stimulated in the presence of competing bacteria, and we tested if biodiversity within bacterial communities affects their antagonistic activity against fungi and fungal-like species. We set up Pseudomonas fluorescens communities of increasing diversity and measured the production of the broad spectrum antifungal compound 2,4-DAPG and their antagonistic activity against different eukaryotes. Diversity increased DAPG concentration and antifungal activity, an effect due to a combination of identity and interactions between species. Our results indicate that investment of pseudomonads into broad spectrum anti-eukaryotic traits is determined by both community composition and diversity and this provides new avenues to understand interactions between bacterial and fungal communities.
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  • 99
    Publication Date: 2014-05-06
    Description: Ecology, Ahead of Print. In some mutualisms cooperation in symbionts is promoted by hosts sanctioning 'cheats' who obtain benefits but fail to reciprocate. In fig-wasp mutualisms agaonid wasps pollinate the trees (Ficus spp.) but are also exploitative by using some flowers as larval food. Ficus can sanction cheats that fail to pollinate by aborting some un-pollinated figs. However, in those un-pollinated figs retained by trees cheats successfully reproduce. When this occurs wasp broods are reduced, suggesting sanctions increase offspring mortality within un-pollinated figs. We investigated sanction mechanisms of abortion and larval mortality against wasp cheats in the monoecious Ficus racemosa, by introducing into figs 1,3,5,7 or 9 female wasps (foundresses) that were either all pollen-laden (P+) or all pollen-free (P-) The abortion rates of P- figs were highest (~60%) when single foundresses were present. Abortion declined with increased foundresses and ceased with seven or more wasps present, irrespective of pollination. In un-aborted figs, wasp fitness (mean offspring per foundress) declined as foundress number increased, especially in P- figs. Reduced broods in P- figs resulted from increased larval mortality of female offspring as foundress number increased, resulting in more male biased sex ratios. Overall sanctions estimated from both abortion rates and reduced offspring production strengthened as the number of cheats increased. In a second experiment we decoupled pollination from wasp oviposition by introducing one pollen-free foundress followed 24h later by seven pollen-laden ovipositor-excised wasps. Compared with P+ and P- single foundress figs, delayed pollination resulted in intermediate larval mortality and wasp fitness, which concurred with patterns of female offspring production. We conclude that fig abortion reflects both pollinator numbers and pollen presence. Sanctions within P- figs initiate soon after oviposition and discriminate against female offspring, thus reducing the benefits to cheats from adaptively biasing their offspring sex ratios. We suggest that costs to cheats via these discriminative sanctions are likely to promote stability in this mutualism.
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  • 100
    Publication Date: 2014-05-06
    Description: Ecology, Ahead of Print. Plants, bacteria and fungi produce essential amino acids (EAAs) with distinctive patterns of δ13C values that can be used as naturally occurring fingerprints of biosynthetic origin of EAAs in a food web. Because animals cannot synthesize EAAs and must obtain them from food, their tissues reflect δ13CEAA patterns found in diet, but it is not known how microbes responsible for hindgut fermentation in some herbivores influence the δ13C values of EAAs in their host's tissues. We examined whether distinctive δ13C fingerprints of hindgut flora are evident in the tissues of green turtles (Chelonia mydas), which are known to be facultative hindgut fermenters. We determined δ13CEAA values in tissues of green turtles foraging herbivorously in neritic habitats of Hawaii and compared them with those from green, olive ridley and loggerhead turtles foraging carnivorously in oceanic environments of the central and south-east Pacific Ocean. Results of multivariate statistical analysis revealed two distinct groups that could be distinguished based on unique δ13CEAA patterns. A three-end member predictive linear discriminant model indicated that δ13CEAA fingerprints existed in the tissues of carnivorous turtles that resembled patterns found in microalgae, which form the base of an oceanic food web, whereas herbivorous turtles derive EAAs from a bacterial or seagrass source. This study demonstrates the capacity for δ13C fingerprinting to establish the biosynthetic origin of EAAs in higher consumers, and that marine turtles foraging on macroalgal diets appear to receive nutritional supplementation from bacterial symbionts in their digestive system.
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