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Translocation experiments with butterflies reveal limits to enhancement of poleward populations under climate change

There is a pressing need to predict how species will change their geographic ranges under climate change. Projections typically assume that temperature is a primary fitness determinant and that populations near the poleward (and upward) range boundary are preadapted to warming. Thus, poleward, peripheral populations will increase with warming, and these increases facilitate poleward range expansions. We tested the assumption that poleward, pe- ripheral populations are enhanced by warming using 2 butterflies (Erynnis propertius and Papilio zelicaon) that co-occur and have contrasting degrees of host specialization and interpopulation genetic differentiation. We performed a reciprocal translocation experiment between central and poleward, peripheral populations in the field and simulated a translocation experiment that included alternate host plants. We found that the performance of both central and peripheral populations of E. propertius were enhanced during the summer months by temperatures characteristic of the range center but that local adaptation of peripheral populations to winter conditions near the range edge could counteract that enhancement. Further, poleward range expansion in this species is prevented by a lack of host plants. In P. zelicaon, the fitness of central and peripheral populations decreased under extreme sum- mer temperatures that occurred in the field at the range center. Performance in this species also was affected by an interaction of temperature and host plant such that host species strongly medi- ated the fitness of peripheral individuals under differing simulated temperatures. Altogether we have evidence that facilitation of poleward range shifts through enhancement of peripheral populations is unlikely in either study species. Lepidoptera 􏰚 range center 􏰚 range expansion 􏰚 range periphery

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Trends and missing parts in the study of movement ecology

Movement is important to all organisms, and accordingly it is addressed in a huge number of papers in the literature. Of nearly 26,000 papers referring to movement, an estimated 34% focused on movement by measuring it or testing hypotheses about it. This enormous amount of information is difficult to review and high- lights the need to assess the collective completeness of movement studies and identify gaps. We surveyed 1,000 randomly selected papers from 496 journals and compared the facets of movement studied with a suggested framework for movement ecology, consisting of internal state (motivation, physiology), motion and navigation capacities, and external factors (both the physical environment and living organisms), and links among these com- ponents. Most studies simply measured and described the move- ment of organisms without reference to ecological or internal factors, and the most frequently studied part of the framework was the link between external factors and motion capacity. Few studies looked at the effects on movement of navigation capacity, or internal state, and those were mainly from vertebrates. For invertebrates and plants most studies were at the population level, whereas more vertebrate studies were conducted at the individual level. Consideration of only population-level averages promul- gates neglect of between-individual variation in movement, po- tentially hindering the study of factors controlling movement. Terminology was found to be inconsistent among taxa and sub- disciplines. The gaps identified in coverage of movement studies highlight research areas that should be addressed to fully under- stand the ecology of movement. dispersal 􏰚 foraging 􏰚 migration 􏰚 navigation 􏰚 physiology

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Measuring the effectiveness of protected area networks in reducing deforestation

Global efforts to reduce tropical deforestation rely heavily on the establishment of protected areas. Measuring the effectiveness of these areas is difficult because the amount of deforestation that would have occurred in the absence of legal protection cannot be directly observed. Conventional methods of evaluating the effectiveness of protected areas can be biased because protection is not randomly assigned and because protection can induce deforesta- tion spillovers (displacement) to neighboring forests. We demon- strate that estimates of effectiveness can be substantially im- proved by controlling for biases along dimensions that are observable, measuring spatial spillovers, and testing the sensitivity of estimates to potential hidden biases. We apply matching meth- ods to evaluate the impact on deforestation of Costa Rica’s re- nowned protected-area system between 1960 and 1997. We find that protection reduced deforestation: approximately 10% of the protected forests would have been deforested had they not been protected. Conventional approaches to evaluating conservation impact, which fail to control for observable covariates correlated with both protection and deforestation, substantially overesti- mate avoided deforestation (by over 65%, based on our estimates). We also find that deforestation spillovers from protected to un- protected forests are negligible. Our conclusions are robust to potential hidden bias, as well as to changes in modeling assump- tions. Our results show that, with appropriate empirical methods, conservation scientists and policy makers can better understand the relationships between human and natural systems and can use this to guide their attempts to protect critical ecosystem services. avoided deforestation 􏰚 conservation policy 􏰚 empirical evaluation 􏰚 spatial spillovers

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The movement ecology and dynamics of plant communities in fragmented landscapes

A conceptual model of movement ecology has recently been advanced to explain all movement by considering the interaction of four elements: internal state, motion capacity, navigation capacities, and external factors. We modified this framework to generate predictions for species richness dynamics of fragmented plant communities and tested them in experimental landscapes across a 7-year time series. We found that two external factors, dispersal vectors and habitat features, affected species coloniza- tion and recolonization in habitat fragments and their effects varied and depended on motion capacity. Bird-dispersed species richness showed connectivity effects that reached an asymptote over time, but no edge effects, whereas wind-dispersed species richness showed steadily accumulating edge and connectivity effects, with no indication of an asymptote. Unassisted species also showed increasing differences caused by connectivity over time, whereas edges had no effect. Our limited use of proxies for movement ecology (e.g., dispersal mode as a proxy for motion capacity) resulted in moderate predictive power for communities and, in some cases, highlighted the importance of a more complete understanding of movement ecology for predicting how landscape conservation actions affect plant community dynamics. corridors 􏰚 dispersal 􏰚 diversity 􏰚 life-history traits 􏰚 species richness

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Multiple movement modes by large herbivores at multiple spatiotemporal scales

Recent theory suggests that animals should switch facultatively among canonical movement modes as a complex function of internal state, landscape characteristics, motion capacity, and navigational capacity. We tested the generality of this paradigm for free-ranging elk (Cervus elaphus) over 5 orders of magnitude in time (minutes to years) and space (meters to 100 km). At the coarsest spatiotemporal scale, elk shifted from a dispersive to a home-ranging phase over the course of 1–3 years after introduc- tion into a novel environment. At intermediate spatiotemporal scales, elk continued to alternate between movement modes. During the dispersive phase, elk alternated between encamped and exploratory modes, possibly linked to changes in motivational goals from foraging to social bonding. During the home-ranging phase, elk movements were characterized by a complex interplay between attraction to preferred habitat types and memory of previous movements across the home-range. At the finest tempo- ral and spatial scale, elk used area-restricted search while brows- ing, interspersed with less sinuous paths when not browsing. Encountering a patch of high-quality food plants triggered the switch from one mode to the next, creating biphasic movement dynamics that were reinforced by local resource heterogeneity. These patterns suggest that multiphasic structure is fundamental to the movement patterns of elk at all temporal and spatial scales tested. elk 􏰚 foraging 􏰚 group formation 􏰚 motivation

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Southern Ocean acidification: A tipping point at 450-ppm atmospheric CO2

Southern Ocean acidification via anthropogenic CO2 uptake is expected to be detrimental to multiple calcifying plankton species by lowering the concentration of carbonate ion (CO32􏱉) to levels where calcium carbonate (both aragonite and calcite) shells begin to dissolve. Natural seasonal variations in carbonate ion concentrations could either hasten or dampen the future onset of this undersaturation of calcium carbonate. We present a large-scale Southern Ocean observational analysis that examines the seasonal magnitude and variability of CO32􏱉 and pH. Our analysis shows an intense wintertime minimum in CO32􏱉 south of the Antarctic Polar Front and when combined with anthropogenic CO2 uptake is likely to induce aragonite undersaturation when atmospheric CO2 levels reach 􏰜450 ppm. Under the IPCC IS92a scenario, Southern Ocean wintertime aragonite undersaturation is projected to occur by the year 2030 and no later than 2038. Some prominent calcifying plankton, in particular the Pteropod species Limacina helicina, have important veliger larval development during winter and will have to experience detrimental carbonate conditions much earlier than previously thought, with possible deleterious flow-on impacts for the wider Southern Ocean marine ecosystem. Our results highlight the critical importance of understanding seasonal carbon dynamics within all calcifying marine ecosystems such as continental shelves and coral reefs, because natural variability may potentially hasten the onset of future ocean acidification. carbon cycle 􏰚 climate change

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Movement ecology of migration in turkey vultures

We develop individual-based movement ecology models (MEM) to explore turkey vulture (Cathartes aura) migration decisions at both hourly and daily scales. Vulture movements in 10 migration events were recorded with satellite-reporting GPS sensors, and flight behavior was observed visually, aided by on-the-ground VHF radio-track- ing. We used the North American Regional Reanalysis dataset to obtain values for wind speed, turbulent kinetic energy (TKE), and cloud height and used a digital elevation model for a measure of terrain ruggedness. A turkey vulture fitted with a heart-rate logger during 124 h of flight during 38 contiguous days showed only a small increase in mean heart rate as distance traveled per day increased, which suggests that, unlike flapping, soaring flight does not lead to greatly increased metabolic costs. Data from 10 migrations for 724 hourly segments and 152 daily segments showed that vultures depended heavily upon high levels of TKE in the atmospheric bound- ary layer to increase flight distances and maintain preferred bearings at both hourly and daily scales. We suggest how the MEM can be extended to other spatial and temporal scales of avian migration. Our success in relating model-derived atmospheric variables to migration indicates the potential of using regional reanalysis data, as here, and potentially other regional, higher-resolution, atmospheric models in predicting changing movement patterns of soaring birds under var- ious scenarios of climate and land use change. energetics 􏰚 flight 􏰚 meteorology

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Understanding strategies for seed dispersal by wind under contrasting atmospheric conditions

Traits associated with seed dispersal vary tremendously among sympatric wind-dispersed plants. We used two contrasting tropical tree species, seed traps, micrometeorology, and a mechanistic model to evaluate how variation in four key traits affects seed dispersal by wind. The conceptual framework of movement ecology, wherein external factors (wind) interact with internal factors (plant traits) that enable movement and determine when and where movement occurs, fully captures the variable inputs and outputs of wind dispersal models and informs their interpretation. We used model calculations to evaluate the spatial pattern of dispersed seeds for the 16 factorial combinations of four traits. The study species differed dramatically in traits related to the timing of seed release, and a strong species by season interaction affected most aspects of the spatial pattern of dispersed seeds. A rich interplay among plant traits and seasonal differences in atmo- spheric conditions caused this interaction. Several of the same plant traits are crucial for both seed dispersal and other aspects of life history variation. Observed traits that limit dispersal are likely to be constrained by their life history consequences. atmospheric turbulence 􏰚 conditional seed release 􏰚 Coupled Eulerian-Lagrangian closure (CELC) model 􏰚 long distance dispersal 􏰚 tropical forest

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Individual movement behavior, matrix heterogeneity, and the dynamics of spatially structured populations

The dynamics of spatially structured populations is characterized by within- and between-patch processes. The available theory describes the latter with simple distance-dependent functions that depend on landscape properties such as interpatch distance or patch size. Despite its potential role, we lack a good mechanistic understanding of how the movement of individuals between patches affects the dynamics of these populations. We used the theoretical framework provided by movement ecology to make a direct representation of the processes determining how individuals connect local populations in a spatially structured population of Iberian lynx. Interpatch processes depended on the heterogeneity of the matrix where patches are embedded and the parameters defining individual movement behavior. They were also very sensitive to the dynamic demographic variables limiting the time moving, the within-patch dynamics of available settlement sites (both spatiotemporally heterogeneous) and the response of indi- viduals to the perceived risk while moving. These context- dependent dynamic factors are an inherent part of the movement process, producing connectivities and dispersal kernels whose variability is affected by other demographic processes. Mechanistic representations of interpatch movements, such as the one pro- vided by the movement-ecology framework, permit the dynamic interaction of birth–death processes and individual movement behavior, thus improving our understanding of stochastic spatially structured populations. demography 􏰚 Iberian lynx 􏰚 metapopulation 􏰚 population dynamics 􏰚 source-sink

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Populations of migratory bird species that did not show a phenological response to climate change are declining

Recent rapid climatic changes are associated with dramatic changes in phenology of plants and animals, with optimal timing of reproduction advancing considerably in the northern hemisphere. Some species may not have advanced their timing of breeding suffi- ciently to continue reproducing optimally relative to the occur- rence of peak food availability, thus becoming mismatched com- pared with their food sources. The degree of mismatch may differ among species, and species with greater mismatch may be char- acterized by declining populations. Here we relate changes in spring migration timing by 100 European bird species since 1960, considered as an index of the phenological response of bird species to recent climate change, to their population trends. Species that declined in the period 1990–2000 did not advance their spring migration, whereas those with stable or increasing populations advanced their migration considerably. On the other hand, popu- lation trends during 1970–1990 were predicted by breeding hab- itat type, northernmost breeding latitude, and winter range (with species of agricultural habitat, breeding at northern latitudes, and wintering in Africa showing an unfavorable conservation status), but not by change in migration timing. The association between population trend in 1990 –2000 and change in migration phenology was not confounded by any of the previously identified predictors of population trends in birds, or by similarity in phenotype among taxa due to common descent. Our findings imply that ecological factors affecting population trends can change over time and suggest that ongoing climatic changes will increasingly threaten vulnerable migratory bird species, augmenting their extinction risk. conservation 􏰚 extinction risk 􏰚 migration phenology 􏰚 population trends

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Sharply increased insect herbivory during the Paleocene–Eocene Thermal Maximum

The Paleocene–Eocene Thermal Maximum (PETM, 55.8 Ma), an abrupt global warming event linked to a transient increase in pCO2, was comparable in rate and magnitude to modern anthropogenic climate change. Here we use plant fossils from the Bighorn Basin of Wyoming to document the combined effects of temperature and pCO2 on insect herbivory. We examined 5,062 fossil leaves from five sites positioned before, during, and after the PETM (59–55.2 Ma). The amount and diversity of insect damage on angiosperm leaves, as well as the relative abundance of specialized damage, correlate with rising and falling temperature. All reach distinct maxima during the PETM, and every PETM plant species is exten- sively damaged and colonized by specialized herbivores. Our study suggests that increased insect herbivory is likely to be a net long-term effect of anthropogenic pCO2 increase and warming temperatures. Bighorn Basin 􏰚 paleobotany 􏰚 plant–insect interactions 􏰚 rapid climate change

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Global declines in oceanic nitrification rates as a consequence of ocean acidification

Ocean acidification produced by dissolution of anthropogenic carbon dioxide (CO2) emissions in seawater has profound conse- quences for marine ecology and biogeochemistry. The oceans have absorbed one-third of CO2 emissions over the past two centuries, altering ocean chemistry, reducing seawater pH, and affecting marine animals and phytoplankton in multiple ways. Microbially mediated ocean biogeochemical processes will be pivotal in deter- mining how the earth system responds to global environmental change; however, how they may be altered by ocean acidification is largely unknown. We show here that microbial nitrification rates decreased in every instance when pH was experimentally reduced (by 0.05–0.14) at multiple locations in the Atlantic and Pacific Oceans. Nitrification is a central process in the nitrogen cycle that produces both the greenhouse gas nitrous oxide and oxidized forms of nitrogen used by phytoplankton and other microorgan- isms in the sea; at the Bermuda Atlantic Time Series and Hawaii Ocean Time-series sites, experimental acidification decreased am- monia oxidation rates by 38% and 36%. Ammonia oxidation rates were also strongly and inversely correlated with pH along a gradi- ent produced in the oligotrophic Sargasso Sea (r2 = 0.87, P < 0.05). Across all experiments, rates declined by 8–38% in low pH treat- ments, and the greatest absolute decrease occurred where rates were highest off the California coast. Collectively our results sug- gest that ocean acidification could reduce nitrification rates by 3–44% within the next few decades, affecting oceanic nitrous oxide production, reducing supplies of oxidized nitrogen in the upper layers of the ocean, and fundamentally altering nitrogen cycling in the sea.

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Insects take a bigger bite out of plants in a warmer, higher carbon dioxide world

Excerpt from text : Because of the direct effect of CO2 and temperature on global food supplies, the influence of these changes on plant physiology and ecology is being actively studied (4–7). How these elements of global change may alter the interactions between plants and the insects that feed on them is relatively unknown. By bringing to light secrets contained in the fossil record, Currano et al. (8), published in this issue of PNAS, found that the amount and diversity of insect damage to plants increased in association with an abrupt rise in atmospheric CO2 and global temperature that occurred 􏰞55 million years ago. If the past is indeed a window to the future, their findings sug- gest that increased insect herbivory will be one more unpleasant surprise arising from anthropogenic climate change.

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Higher origination and extinction rates in larger mammals

Do large mammals evolve faster than small mammals or vice versa? Because the answer to this question contributes to our understanding of how life-history affects long-term and large-scale evolutionary patterns, and how microevolutionary rates scale-up to macroevolu- tionary rates, it has received much attention. A satisfactory or con- sistent answer to this question is lacking, however. Here, we take a fresh look at this problem using a large fossil dataset of mammals from the Neogene of the Old World (NOW). Controlling for sampling biases, calculating per capita origination and extinction rates of boundary-crossers and estimating survival probabilities using cap- ture-mark-recapture (CMR) methods, we found the recurring pattern that large mammal genera and species have higher origination and extinction rates, and therefore shorter durations. This pattern is surprising in the light of molecular studies, which show that smaller animals, with their shorter generation times and higher metabolic rates, have greater absolute rates of evolution. However, higher molecular rates do not necessarily translate to higher taxon rates because both the biotic and physical environments interact with phenotypic variation, in part fueled by mutations, to affect origina- tion and extinction rates. To explain the observed pattern, we propose that the ability to evolve and maintain behavior such as hibernation, torpor and burrowing, collectively termed ‘‘sleep-or- hide’’ (SLOH) behavior, serves as a means of environmental buffering during expected and unexpected environmental change. SLOH be- havior is more common in some small mammals, and, as a result, SLOH small mammals contribute to higher average survivorship and lower origination probabilities among small mammals. body size 􏰚 environmental buffering 􏰚 metabolism 􏰚 Neogene mammals 􏰚 turnover

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Higher effect of plant species diversity on productivity in natural than artificial ecosystems

Current and expected changes in biodiversity have motivated major experiments, which reported a positive relationship be- tween plant species diversity and primary production. As a first step in addressing this relationship, these manipulative experi- ments controlled as many potential confounding covariables as possible and assembled artificial ecosystems for the purpose of the experiments. As a new step in this endeavor, we asked how plant species richness relates to productivity in a natural ecosystem. Here, we report on an experiment conducted in a natural ecosys- tem in the Patagonian steppe, in which we assessed the biodiver- sity effect on primary production. Using a plant species diversity gradient generated by removing species while maintaining con- stant biomass, we found that aboveground net primary production increased with the number of plant species. We also found that the biodiversity effect was larger in natural than in artificial ecosys- tems. This result supports previous findings and also suggests that the effect of biodiversity in natural ecosystems may be much larger than currently thought. biodiversity 􏰚 carbon cycle 􏰚 ecosystem functioning 􏰚 Patagonian steppe 􏰚 resource partitioning

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The impact of Miocene atmospheric carbon dioxide fluctuations on climate and the evolution of terrestrial ecosystems

The Miocene is characterized by a series of key climatic events that led to the founding of the late Cenozoic icehouse mode and the dawn of modern biota. The processes that caused these developments, and particularly the role of atmospheric CO2 as a forcing factor, are poorly understood. Here we present a CO2 record based on stomatal frequency data from multiple tree species. Our data show striking CO2 fluctuations of 􏰜600–300 parts per million by volume (ppmv). Periods of low CO2 are contemporaneous with major glaciations, whereas elevated CO2 of 500 ppmv coincides with the climatic optimum in the Miocene. Our data point to a long-term coupling between atmospheric CO2 and climate. Major changes in Miocene terrestrial ecosystems, such as the expansion of grasslands and radiations among terrestrial herbivores such as horses, can be linked to these marked fluctuations in CO2. atmospheric CO2 􏰚 fossil plants 􏰚 paleoclimates 􏰚 stomata 􏰚 C4 plants

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Physiology and Climate Change

Studies of physiological mechanisms are needed to predict climate effects on ecosystems at species and community levels.

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Future hotspots of terrestrial mammal loss

Current levels of endangerment and historical trends of species and habitats are the main criteria used to direct conservation efforts globally. Estimates of future declines, which might indicate different priorities than past declines, have been limited by the lack of appropriate data and models. Given that much of con- servation is about anticipating and responding to future threats, our inability to look forward at a global scale has been a major constraint on effective action. Here, we assess the geography and extent of projected future changes in suitable habitat for terrestrial mammals within their present ranges. We used a global earth-system model, IMAGE, coupled with fine-scale habitat suitability models and parametrized accord- ing to four global scenarios of human development. We identified the most affected countries by 2050 for each scenario, assuming that no additional conservation actions other than those described in the scenarios take place. We found that, with some exceptions, most of the countries with the largest predicted losses of suitable habitat for mammals are in Africa and the Americas. African and North American countries were also predicted to host the most species with large proportional global declines. Most of the countries we identified as future hotspots of terrestrial mammal loss have little or no overlap with the present global conservation priorities, thus confirming the need for forward-looking analyses in conservation priority setting. The expected growth in human populations and consumption in hotspots of future mammal loss mean that local conservation actions such as protected areas might not be sufficient to mitigate losses. Other policies, directed towards the root causes of biodiversity loss, are required, both in Africa and other parts of the world.

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Traversing the mountaintop: world fossil fuel production to 2050

During the past century, fossil fuels—petroleum liquids, natural gas and coal—were the dominant source of world energy production. From 1950 to 2005, fossil fuels provided 85–93% of all energy production. All fossil fuels grew substantially during this period, their combined growth exceeding the increase in world population. This growth, however, was irregular, providing for rapidly grow- ing per capita production from 1950 to 1980, stable per capita production from 1980 to 2000 and rising per capita production again after 2000. During the past half century, growth in fossil fuel pro- duction was essentially limited by energy demand. During the next half century, fossil fuel production will be limited primarily by the amount and characteristics of remaining fossil fuel resources. Three possible scenarios—low, medium and high—are developed for the production of each of the fossil fuels to 2050. These scenarios differ primarily by the amount of ultimate resources estimated for each fossil fuel. Total fossil fuel production will continue to grow, but only slowly for the next 15–30 years. The subsequent peak plateau will last for 10–15 years. These production peaks are robust; none of the fossil fuels, even with highly optimistic resource estimates, is projected to keep growing beyond 2050. World fossil fuel production per capita will thus begin an irreversible decline between 2020 and 2030. Keywords: coal; fossil fuels; natural gas; peak fuel production; petroleum liquids; production scenarios

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Increasing River Discharge to the Arctic Ocean

Synthesis of river-monitoring data reveals that the average annual discharge of fresh water from the six largest Eurasian rivers to the Arctic Ocean increased by7%from1936to1999.Theaverageannualrateofincreasewas2.0􏰤0.7 cubic kilometers per year. Consequently, average annual discharge from the six rivers is now about 128 cubic kilometers per year greater than it was when routine measurements of discharge began. Discharge was correlated with changes in both the North Atlantic Oscillation and global mean surface air temperature. The observed large-scale change in freshwater flux has potentially important implications for ocean circulation and climate.

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