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Prey choice and habitat use drive sea otter pathogen exposure in a resource-limited coastal system

The processes promoting disease in wild animal populations are highly complex, yet identifying these processes is critically important for conservation when disease is limiting a population. By combining field studies with epidemiologic tools, we evaluated the relationship between key factors impeding southern sea otter ( Enhydra lutris nereis ) population growth: disease and resource limitation. This threatened population has struggled to recover despite protection, so we followed radio-tagged sea otters and evaluated infection with 2 disease-causing protozoal pathogens, Toxoplasma gondii and Sarcocystis neurona , to reveal risks that increased the likelihood of pathogen exposure. We identified patterns of pathogen infection that are linked to individual animal behavior, prey choice, and habitat use. We detected a high-risk spatial cluster of S. neurona infections in otters with home ranges in southern Monterey Bay and a coastal segment near San Simeon and Cambria where otters had high levels of infection with T. gondii . We found that otters feeding on abalone, which is the preferred prey in a resource-abundant marine ecosystem, had a very low risk of infection with either pathogen, whereas otters consuming small marine snails were more likely to be infected with T. gondii . Individual dietary specialization in sea otters is an adaptive mechanism for coping with limited food resources along central coastal California. High levels of infection with protozoal pathogens may be an adverse consequence of dietary specialization in this threatened species, with both depleted resources and disease working synergistically to limit recovery.

California

Elevated CO 2 stimulates marsh elevation gain, counterbalancing sea-level rise

Tidal wetlands experiencing increased rates of sea-level rise (SLR) must increase rates of soil elevation gain to avoid permanent conversion to open water. The maximal rate of SLR that these ecosystems can tolerate depends partly on mineral sediment deposition, but the accumulation of organic matter is equally important for many wetlands. Plant productivity drives organic matter dynamics and is sensitive to global change factors, such as rising atmospheric CO 2 concentration. It remains unknown how global change will influence organic mechanisms that determine future tidal wetland viability. Here, we present experimental evidence that plant response to elevated atmospheric [CO 2 ] stimulates biogenic mechanisms of elevation gain in a brackish marsh. Elevated CO 2 (ambient + 340 ppm) accelerated soil elevation gain by 3.9 mm yr −1 in this 2-year field study, an effect mediated by stimulation of below-ground plant productivity. Further, a companion greenhouse experiment revealed that the CO 2 effect was enhanced under salinity and flooding conditions likely to accompany future SLR. Our results indicate that by stimulating biogenic contributions to marsh elevation, increases in the greenhouse gas, CO 2 , may paradoxically aid some coastal wetlands in counterbalancing rising seas.

Proceedings of the National Academy of Sciences

Sexual selection affects local extinction and turnover in bird communities

Predicting extinction risks has become a central goal for conservation and evolutionary biologists interested in population and community dynamics. Several factors have been put forward to explain risks of extinction, including ecological and life history characteristics of individuals. For instance, factors that affect the balance between natality and mortality can have profound effects on population persistence. Sexual selection has been identified as one such factor. Populations under strong sexual selection experience a number of costs ranging from increased predation and parasitism to enhanced sensitivity to environmental and demographic stochasticity. These findings have led to the prediction that local extinction rates should be higher for species/populations with intense sexual selection. We tested this prediction by analyzing the dynamics of natural bird communities at a continental scale over a period of 21 years (1975-1996), using relevant statistical tools. In agreement with the theoretical prediction, we found that sexual selection increased risks of local extinction (dichromatic birds had on average a 23% higher local extinction rate than monochromatic species). However, despite higher local extinction probabilities, the number of dichromatic species did not decrease over the period considered in this study. This pattern was caused by higher local turnover rates of dichromatic species, resulting in relatively stable communities for both groups of species. Our results suggest that these communities function as metacommunities, with frequent local extinctions followed by colonization. Anthropogenic factors impeding dispersal might therefore have a significant impact on the global persistence of sexually selected species.

Proceedings of the National Academy of Sciences

Quantifying ecological, morphological, and genetic variation to delimit species in the coast horned lizard species complex (Phrynosoma)

Lineage separation and divergence form a temporally extended process whereby populations may diverge genetically, morphologically, or ecologically, and these contingent properties of species provide the operational criteria necessary for species delimitation.We inferred the historical process of lineage formation in the coast horned lizard (Phrynosoma coronatum) species complex by evaluating a diversity of operational species criteria, including divergence in mtDNA (98 specimens; 2,781 bp) and nuclear loci (RAG-1, 1,054 bp; BDNF 529 bp), ecological niches (11 bioclimatic variables; 285 unique localities), and cranial horn shapes (493 specimens; 16 landmarks). A phylogenetic analysis of mtDNA recovers 5 phylogeographic groups arranged latitudinally along the Baja California Peninsula and in California. The 2 southern phylogeographic groups exhibit concordance between genetic, morphological, and ecological divergence; however, differentiation is weak or absent at more recent levels defined by phylogeographic breaks in California. Interpreting these operational species criteria together suggests that there are 3 ecologically divergent and morphologically diagnosable species within the P. coronatum complex. Our 3-species taxonomic hypothesis invokes a deep coalescence event when fitting the mtDNA genealogy into the species tree, which is not unexpected for populations that have diverged recently. Although the hypothesis that the 3 phylogeographic groups distributed across California each represent distinctive species is not supported by all of the operational species criteria evaluated in this study, the conservation status of the imperiled populations represented by these genealogical units remains critical.

Proceedings of the National Academy of Sciences of

Temporal latitudinal-gradient dynamics and tropical instability of deep-sea species diversity

A benthic microfaunal record from the equatorial Atlantic Ocean over the past four glacial-interglacial cycles was investigated to understand temporal dynamics of deep-sea latitudinal species diversity gradients (LSDGs). The results demonstrate unexpected instability and high amplitude fluctuations of species diversity in the tropical deep ocean that are correlated with orbital-scale oscillations in global climate: Species diversity is low during glacial and high during interglacial periods. This implies that climate severely influences deep-sea diversity, even at tropical latitudes, and that deep-sea LSDGs, while generally present for the last 36 million years, were weakened or absent during glacial periods. Temporally dynamic LSDGs and unstable tropical diversity require reconsideration of current ecological hypotheses about the generation and maintenance of biodiversity as they apply to the deep sea, and underscore the potential vulnerability and conservation importance of tropical deep-sea ecosystems.

Proceedings of the National Academy of Sciences of

Housing growth in and near United States protected areas limits their conservation value

Protected areas are crucial for biodiversity conservation because they provide safe havens for species threatened by land-use change and resulting habitat loss. However, protected areas are only effective when they stop habitat loss within their boundaries, and are connected via corridors to other wild areas. The effectiveness of protected areas is threatened by development; however, the extent of this threat is unknown. We compiled spatially-detailed housing growth data from 1940 to 2030, and quantified growth for each wilderness area, national park, and national forest in the conterminous United States. Our findings show that housing development in the United States may severely limit the ability of protected areas to function as a modern "Noah's Ark." Between 1940 and 2000, 28 million housing units were built within 50 km of protected areas, and 940,000 were built within national forests. Housing growth rates during the 1990s within 1 km of protected areas (20% per decade) outpaced the national average (13%). If long-term trends continue, another 17 million housing units will be built within 50 km of protected areas by 2030 (1 million within 1 km), greatly diminishing their conservation value. US protected areas are increasingly isolated, housing development in their surroundings is decreasing their effective size, and national forests are even threatened by habitat loss within their administrative boundaries. Protected areas in the United States are thus threatened similarly to those in developing countries. However, housing growth poses the main threat to protected areas in the United States whereas deforestation is the main threat in developing countries.

Proceedings of the National Academy of Sciences of

Response of Colorado river runoff to dust radiative forcing in snow

The waters of the Colorado River serve 27 million people in seven states and two countries but are overallocated by more than 10% of the river's historical mean. Climate models project runoff losses of 7-20% from the basin in this century due to human-induced climate change. Recent work has shown however that by the late 1800s, decades prior to allocation of the river's runoff in the 1920s, a fivefold increase in dust loading from anthropogenically disturbed soils in the southwest United States was already decreasing snow albedo and shortening the duration of snow cover by several weeks. The degree to which this increase in radiative forcing by dust in snow has affected timing and magnitude of runoff from the Upper Colorado River Basin (UCRB) is unknown. Hereweuse the Variable Infiltration Capacity model with postdisturbance and predisturbance impacts of dust on albedo to estimate the impact on runoff from the UCRB across 1916-2003. We find that peak runoff at Lees Ferry, Arizona has occurred on average 3 wk earlier under heavier dust loading and that increases in evapotranspiration from earlier exposure of vegetation and soils decreases annual runoff by more than 1.0 billion cubic meters or ???5% of the annual average. The potential to reduce dust loading through surface stabilization in the deserts and restore more persistent snow cover, slow runoff, and increase water resources in the UCRB may represent an important mitigation opportunity to reduce system management tensions and regional impacts of climate change.

Proceedings of the National Academy of Sciences of

11-Deoxycortisol is a corticosteroid hormone in the lamprey

Corticosteroid hormones are critical for controlling metabolism, hydromineral balance, and the stress response in vertebrates. Although corticosteroid hormones have been well characterized in most vertebrate groups, the identity of the earliest vertebrate corticosteroid hormone has remained elusive. Here we provide evidence that 11-deoxycortisol is the corticosteroid hormone in the lamprey, a member of the agnathans that evolved more than 500 million years ago. We used RIA, HPLC, and mass spectrometry analysis to determine that 11-deoxycortisol is the active corticosteroid present in lamprey plasma. We also characterized an 11-deoxycortisol receptor extracted from sea lamprey gill cytosol. The receptor was highly specific for 11-deoxycortisol and exhibited corticosteroid binding characteristics, including DNA binding. Furthermore, we observed that 11-deoxycortisol was regulated by the hypothalamus-pituitary axis and responded to acute stress. 11-Deoxycortisol implants reduced sex steroid concentrations and upregulated gill Na+, K+-ATPase, an enzyme critical for ion balance. We show here that 11-deoxycortisol functioned as both a glucocorticoid and a mineralocorticoid in the lamprey. Our findings indicate that a complex and highly specific corticosteroid signaling pathway evolved at least 500 million years ago with the arrival of the earliest vertebrate.

Proceedings of the National Academy of Sciences of

Use of multiple dispersal pathways facilitates amphibian persistence in stream networks

Although populations of amphibians are declining worldwide, there is no evidence that salamanders occupying small streams are experiencing enigmatic declines, and populations of these species seem stable. Theory predicts that dispersal through multiple pathways can stabilize populations, preventing extinction in habitat networks. However, empirical data to support this prediction are absent for most species, especially those at risk of decline. Our mark-recapture study of stream salamanders reveals both a strong upstream bias in dispersal and a surprisingly high rate of overland dispersal to adjacent headwater streams. This evidence of route-dependent variation in dispersal rates suggests a spatial mechanism for population stability in headwater-stream salamanders. Our results link the movement behavior of stream salamanders to network topology, and they underscore the importance of identifying and protecting critical dispersal pathways when addressing region-wide population declines.

Proceedings of the National Academy of Sciences of

Long-term reductions in anthropogenic nutrients link to improvements in Chesapeake Bay habitat

Great effort continues to focus on ecosystem restoration and reduction of nutrient inputs thought to be responsible, in part, for declines in estuary habitats worldwide. The ability of environmental policy to address restoration is limited, in part, by uncertainty in the relationships between costly restoration and benefits. Here, we present results from an 18-y field investigation (1990-2007) of submerged aquatic vegetation (SAV) community dynamics and water quality in the Potomac River, a major tributary of the Chesapeake Bay. River and anthropogenic discharges lower water clarity by introducing nutrients that stimulate phytoplankton and epiphyte growth as well as suspended sediments. Efforts to restore the Chesapeake Bay are often viewed as failing. Overall nutrient reduction and SAV restoration goals have not been met. In the Potomac River, however, reduced in situ nutrients, wastewater-treatment effluent nitrogen, and total suspended solids were significantly correlated to increased SAV abundance and diversity. Species composition and relative abundance also correlated with nutrient and water-quality conditions, indicating declining fitness of exotic species relative to native species during restoration. Our results suggest that environmental policies that reduce anthropogenic nutrient inputs do result in improved habitat quality, with increased diversity and native species abundances. The results also help elucidate why SAV cover has improved only in some areas of the Chesapeake Bay.

Chesapeake Bay

Ecological contingency in the effects of climatic warming on forest herb communities

Downscaling from the predictions of general climate models is critical to current strategies for mitigating species loss caused by climate change. A key impediment to this downscaling is that we lack a fully developed understanding of how variation in physical, biological, or land-use characteristics mediates the effects of climate change on ecological communities within regions. We analyzed change in understory herb communities over a 60-y period (1949/1951–2007/2009) in a complex montane landscape (the Siskiyou Mountains, Oregon) where mean temperatures have increased 2 °C since 1948, similar to projections for other terrestrial communities. Our 185 sites included primary and secondary-growth lower montane forests (500–1.200 m above sea level) and primary upper montane to subalpine forests (1,500–2,100 m above sea level). In lower montane forests, regardless of land-use history, we found multiple herb-community changes consistent with an effectively drier climate, including lower mean specific leaf area, lower relative cover by species of northern biogeographic affinity, and greater compositional resemblance to communities in southerly topographic positions. At higher elevations we found qualitatively different and more modest changes, including increases in herbs of northern biogeographic affinity and in forest canopy cover. Our results provide community-level validation of predicted nonlinearities in climate change effects.

Oregon

Reclaiming freshwater sustainability in the Cadillac Desert

Increasing human appropriation of freshwater resources presents a tangible limit to the sustainability of cities, agriculture, and ecosystems in the western United States. Marc Reisner tackles this theme in his 1986 classic Cadillac Desert: The American West and Its Disappearing Water . Reisner's analysis paints a portrait of region-wide hydrologic dysfunction in the western United States, suggesting that the storage capacity of reservoirs will be impaired by sediment infilling, croplands will be rendered infertile by salt, and water scarcity will pit growing desert cities against agribusiness in the face of dwindling water resources. Here we evaluate these claims using the best available data and scientific tools. Our analysis provides strong scientific support for many of Reisner's claims, except the notion that reservoir storage is imminently threatened by sediment. More broadly, we estimate that the equivalent of nearly 76% of streamflow in the Cadillac Desert region is currently appropriated by humans, and this figure could rise to nearly 86% under a doubling of the region's population. Thus, Reisner's incisive journalism led him to the same conclusions as those rendered by copious data, modern scientific tools, and the application of a more genuine scientific method. We close with a prospectus for reclaiming freshwater sustainability in the Cadillac Desert, including a suite of recommendations for reducing region-wide human appropriation of streamflow to a target level of 60%.

Proceedings of the National Academy of Sciences of

Host and viral ecology determine bat rabies seasonality and maintenance

Rabies is an acute viral infection that is typically fatal. Most rabies modeling has focused on disease dynamics and control within terrestrial mammals (e.g., raccoons and foxes). As such, rabies in bats has been largely neglected until recently. Because bats have been implicated as natural reservoirs for several emerging zoonotic viruses, including SARS-like corona viruses, henipaviruses, and lyssaviruses, understanding how pathogens are maintained within a population becomes vital. Unfortunately, little is known about maintenance mechanisms for any pathogen in bat populations. We present a mathematical model parameterized with unique data from an extensive study of rabies in a Colorado population of big brown bats (Eptesicus fuscus) to elucidate general maintenance mechanisms. We propose that life history patterns of many species of temperate-zone bats, coupled with sufficiently long incubation periods, allows for rabies virus maintenance. Seasonal variability in bat mortality rates, specifically low mortality during hibernation, allows long-term bat population viability. Within viable bat populations, sufficiently long incubation periods allow enough infected individuals to enter hibernation and survive until the following year, and hence avoid an epizootic fadeout of rabies virus. We hypothesize that the slowing effects of hibernation on metabolic and viral activity maintains infected individuals and their pathogens until susceptibles from the annual birth pulse become infected and continue the cycle. This research provides a context to explore similar host ecology and viral dynamics that may explain seasonal patterns and maintenance of other bat-borne diseases.

Proceedings of the National Academy of Sciences of

Ecosystem impacts of three sequential hurricanes (Dennis, Floyd, and Irene) on the United States' largest lagoonal estuary, Pamlico Sound, NC

Three sequential hurricanes, Dennis, Floyd, and Irene, affected coastal North Carolina in September and October 1999. These hurricanes inundated the region with up to 1 m of rainfall, causing 50- to 500-year flooding in the watershed of the Pamlico Sound, the largest lagoonal estuary in the United States and a key West Atlantic fisheries nursery. We investigated the ecosystem-level impacts on and responses of the Sound to the floodwater discharge. Floodwaters displaced three-fourths of the volume of the Sound, depressed salinity by a similar amount, and delivered at least half of the typical annual nitrogen load to this nitrogen-sensitive ecosystem. Organic carbon concentrations in floodwaters entering Pamlico Sound via a major tributary (the Neuse River Estuary) were at least 2-fold higher than concentrations under prefloodwater conditions. A cascading set of physical, chemical, and ecological impacts followed, including strong vertical stratification, bottom water hypoxia, a sustained increase in algal biomass, displacement of many marine organisms, and a rise in fish disease. Because of the Sound's long residence time (???1 year), we hypothesize that the effects of the short-term nutrient enrichment could prove to be multiannual. A predicted increase in the frequency of hurricane activity over the next few decades may cause longer-term biogeochemical and trophic changes in this and other estuarine and coastal habitats.

Proceedings of the National Academy of Sciences of

Large carbon isotope fractionation associated with oxidation of methyl halides by methylotrophic bacteria

The largest biological fractionations of stable carbon isotopes observed in nature occur during production of methane by methanogenic archaea. These fractionations result in substantial (as much as ≈70‰) shifts in δ 13 C relative to the initial substrate. We now report that a stable carbon isotopic fractionation of comparable magnitude (up to 70‰) occurs during oxidation of methyl halides by methylotrophic bacteria. We have demonstrated biological fractionation with whole cells of three methylotrophs (strain IMB-1, strain CC495, and strain MB2) and, to a lesser extent, with the purified cobalamin-dependent methyltransferase enzyme obtained from strain CC495. Thus, the genetic similarities recently reported between methylotrophs, and methanogens with respect to their pathways for C 1 -unit metabolism are also reflected in the carbon isotopic fractionations achieved by these organisms. We found that only part of the observed fractionation of carbon isotopes could be accounted for by the activity of the corrinoid methyltransferase enzyme, suggesting fractionation by enzymes further along the degradation pathway. These observations are of potential biogeochemical significance in the application of stable carbon isotope ratios to constrain the tropospheric budgets for the ozone-depleting halocarbons, methyl bromide and methyl chloride. Methyl bromide (MeBr) and methyl chloride (MeCl) are, respectively, the most abundant volatile bromo- and chlorocarbons in the troposphere and are major contributors to stratospheric ozone destruction (1). Both compounds have natural and human-influenced sources and a predominant sink by reaction with OH in the troposphere (2–4). MeBr also has a bacterial soil sink (5) that represents about 20% of the estimated total removal from the troposphere, and it is likely that a soil sink of similar magnitude exists for MeCl (6). Hence, if an isotopic fractionation is associated with the soil sink, it will influence the isotopic compositions of MeBr and MeCl in the lower atmosphere (7). The δ 13 C value of industrially produced MeBr ranges between −43.5‰ and −66.4‰ (7), but δ 13 C values of tropospheric MeBr and natural sources are not yet known. The δ 13 C of atmospheric MeCl has been measured from −22‰ to −45‰ (8, 9). If carbon isotope ratios are to be used to constrain the budgets of these methyl halides, it is essential to determine the extent of carbon isotope fractionation that occurs during biological degradation of these compounds. Methylotrophic bacteria use C 1 compounds, which are simple organic molecules that contain no carbon–carbon bonds. Strains IMB-1, CC495, and MB2 are as-yet-unnamed facultative methylotrophs isolated from agricultural soil, woodland leaf litter, and coastal seawater, respectively (10–13), environments where methyl halides are produced. They are members of the α subgroup of the Proteobacteria . On the basis of 16S rRNA gene sequences, strains IMB-1 and CC495 show some phylogenetic alignment with the genus Rhizobium (10, 11) and are very closely related to the new genus Pseudoaminobacte r (I. McDonald, personal communication). Strain MB2 aligns within the Ruegeria clade [J. K. Schaefer, K. D. Goodwin, I. R. McDonald, J. C. Murrell and R.S.O., unpublished work]. All of these aerobic bacteria are methylotrophs in that they can grow by using MeBr or MeCl as their sole carbon source, but they do not metabolize methane. They oxidize MeBr, MeCl, and methyl iodide (MeI) to CO 2 . Soil bacteria are known to consume MeBr at the ambient tropospheric mixing ratio of around 10 parts per trillion by volume (5). Preliminary experiments with strain IMB-1 indicate that it can oxidize MeBr at these mixing ratios** and is therefore likely to be characteristic of bacteria associated with MeBr uptake by soils. We examined δ 13 C of MeCl, MeBr, and MeI during oxidation by whole-cell suspensions of IMB-1 and CC495 and also the change in δ 13 C values of the three methyl halides during oxidation by the marine strain MB2. In addition, we measured the fractionation of carbon isotopes during formation of methane thiol (MeSH) from MeCl by the purified cobalamin-dependent enzyme, halomethane:bisulphide/halide ion methyltransferase (11) from CC495, to determine whether this initial step in MeCl degradation could account for the observed fractionation by whole cells. Finally, we determined the fractionation associated with the degradation of MeBr during field studies with agricultural soil by monitoring MeBr concentration and δ 13 C of MeBr in the headspace of flux chambers under fumigation conditions.

Proceedings of the National Academy of Sciences of

Nitrous oxide emission from denitrification in stream and river networks

Nitrous oxide (N 2 O) is a potent greenhouse gas that contributes to climate change and stratospheric ozone destruction. Anthropogenic nitrogen (N) loading to river networks is a potentially important source of N 2 O via microbial denitrification that converts N to N 2 O and dinitrogen (N 2 ). The fraction of denitrified N that escapes as N 2 O rather than N 2 (i.e., the N 2 O yield) is an important determinant of how much N 2 O is produced by river networks, but little is known about the N 2 O yield in flowing waters. Here, we present the results of whole-stream 15 N-tracer additions conducted in 72 headwater streams draining multiple land-use types across the United States. We found that stream denitrification produces N 2 O at rates that increase with stream water nitrate (NO 3 − ) concentrations, but that <1% of denitrified N is converted to N 2 O. Unlike some previous studies, we found no relationship between the N 2 O yield and stream water NO 3 − . We suggest that increased stream NO 3 − loading stimulates denitrification and concomitant N 2 O production, but does not increase the N 2 O yield. In our study, most streams were sources of N 2 O to the atmosphere and the highest emission rates were observed in streams draining urban basins. Using a global river network model, we estimate that microbial N transformations (e.g., denitrification and nitrification) convert at least 0.68 Tg·y −1 of anthropogenic N inputs to N 2 O in river networks, equivalent to 10% of the global anthropogenic N 2 O emission rate. This estimate of stream and river N 2 O emissions is three times greater than estimated by the Intergovernmental Panel on Climate Change. Humans have more than doubled the availability of fixed nitrogen (N) in the biosphere, particularly through the production of N fertilizers and the cultivation of N-fixing crops ( 1 ). Increasing N availability is producing unintended environmental consequences including enhanced emissions of nitrous oxide (N 2 O), a potent greenhouse gas ( 2 ) and an important cause of stratospheric ozone destruction ( 3 ). The Intergovernmental Panel on Climate Change (IPCC) estimates that the microbial conversion of agriculturally derived N to N 2 O in soils and aquatic ecosystems is the largest source of anthropogenic N 2 O to the atmosphere ( 2 ). The production of N 2 O in agricultural soils has been the focus of intense investigation (i.e., >1,000 published studies) and is a relatively well constrained component of the N 2 O budget ( 4 ). However, emissions of anthropogenic N 2 O from streams, rivers, and estuaries have received much less attention and remain a major source of uncertainty in the global anthropogenic N 2 O budget. Microbial denitrification is a large source of N 2 O emissions in terrestrial and aquatic ecosystems. Most microbial denitrification is a form of anaerobic respiration in which nitrate (NO 3 − , the dominant form of inorganic N) is converted to dinitrogen (N 2 ) and N 2 O gases ( 5 ). The proportion of denitrified NO 3 − that is converted to N 2 O rather than N 2 (hereafter referred to as the N 2 O yield and expressed as the mole ratio) partially controls how much N 2 O is produced via denitrification ( 6 ), but few studies provide information on the N 2 O yield in streams and rivers because of the difficulty of measuring N 2 and N 2 O production in these systems. Here we report rates of N 2 and N 2 O production via denitrification measured using whole-stream 15 NO 3 − -tracer experiments in 72 headwater streams draining different land-use types across the United States. This project, known as the second Lotic Intersite Nitrogen eXperiment (LINX II), provides unique whole-system measurements of the N 2 O yield in streams. Although N 2 O emission rates have been reported for streams and rivers ( 7 , 8 ), the N 2 O yield has been studied mostly in lentic freshwater and marine ecosystems, where it generally ranges between 0.1 and 1.0%, although yields as high as 6% have been observed ( 9 ). These N 2 O yields are low compared with observations in soils (0–100%) ( 10 ), which may be a result of the relatively lower oxygen (O 2 ) availability in the sediments of lakes and estuaries. However, dissolved O 2 in headwater streams is commonly near atmospheric equilibrium and benthic algal biofilms can produce O 2 at the sediment–water interface, resulting in strong redox gradients more akin to those in partially wetted soils. Thus, streams may have variable and often high N 2 O yields, similar to those in soils ( 11 ). The N 2 O yield in headwater streams is of particular interest because much of the NO 3 − input to rivers is derived from groundwater upwelling into headwater streams. Furthermore, headwater streams compose the majority of stream length within a drainage network and have high ratios of bioreactive benthic surface area to water volume ( 12 ).

Proceedings of the National Academy of Sciences of

Fish as major carbonate mud producers and missing components of the tropical carbonate factory

Carbonate mud is a major constituent of recent marine carbonate sediments and of ancient limestones, which contain unique records of changes in ocean chemistry and climate shifts in the geological past. However, the origin of carbonate mud is controversial and often problematic to resolve. Here we show that tropical marine fish produce and excrete various forms of precipitated (nonskeletal) calcium carbonate from their guts (“low” and “high” Mg-calcite and aragonite), but that very fine-grained (mostly < 2 μm) high Mg-calcite crystallites (i.e., MgCO 3 ) are their dominant excretory product. Crystallites from fish are morphologically diverse and species-specific, but all are unique relative to previously known biogenic and abiotic sources of carbonate within open marine systems. Using site specific fish biomass and carbonate excretion rate data we estimate that fish produce ∼6.1 × 10 6 kg CaCO 3 /year across the Bahamian archipelago, all as mud-grade (the < 63 μm fraction) carbonate and thus as a potential sediment constituent. Estimated contributions from fish to total carbonate mud production average ∼14% overall, and exceed 70% in specific habitats. Critically, we also document the widespread presence of these distinctive fish-derived carbonates in the finest sediment fractions from all habitat types in the Bahamas, demonstrating that these carbonates have direct relevance to contemporary carbonate sediment budgets. Fish thus represent a hitherto unrecognized but significant source of fine-grained carbonate sediment, the discovery of which has direct application to the conceptual ideas of how marine carbonate factories function both today and in the past.

Proceedings of the National Academy of Sciences of

The trans-Himalayan flights of bar-headed geese (Anser indicus)

Birds that fly over mountain barriers must be capable of meeting the increased energetic cost of climbing in low-density air, even though less oxygen may be available to support their metabolism. This challenge is magnified by the reduction in maximum sustained climbing rates in large birds. Bar-headed geese ( Anser indicus ) make one of the highest and most iconic transmountain migrations in the world. We show that those populations of geese that winter at sea level in India are capable of passing over the Himalayas in 1 d, typically climbing between 4,000 and 6,000 m in 7–8 h. Surprisingly, these birds do not rely on the assistance of upslope tailwinds that usually occur during the day and can support minimum climb rates of 0.8–2.2 km·h −1 , even in the relative stillness of the night. They appear to strategically avoid higher speed winds during the afternoon, thus maximizing safety and control during flight. It would seem, therefore, that bar-headed geese are capable of sustained climbing flight over the passes of the Himalaya under their own aerobic power.

Himalayas