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Article ; Online: Modeling Patterns and Controls of Food Web Structure in Saline Wetlands of a Rocky Mountain Basin

Lovvorn, James R.

Ecosystems. 2023 Mar., v. 26, no. 2 p.457-472

2023

Abstract: Semiarid intermountain basins of western North America contain many wetlands driven by flood irrigation from mountain snowpack. With decreasing snowpack and increasing urban water demands, maintaining these islands of productivity within otherwise dry ... ...

Abstract	Semiarid intermountain basins of western North America contain many wetlands driven by flood irrigation from mountain snowpack. With decreasing snowpack and increasing urban water demands, maintaining these islands of productivity within otherwise dry shortgrass steppe requires understanding factors that support wetland food webs. In the Laramie Basin, Wyoming, USA the two main wetland types differ in consistency of freshwater inflow and salinity—oligosaline (0.5–5‰ TDS) wetlands dominated by the macroalgae Chara spp., and mesosaline (5–18‰) wetlands dominated by the submersed vascular plant Potamogeton pectinatus. Food web models indicate that production of epiphytic and benthic microalgae and bacteria was limiting to higher trophic levels. Production of benthic microalgae in more shaded sediments within Chara stands was only 11% of that in Potamogeton stands. However, total microalgal production was 32% greater in Chara stands because epiphyton production was 3 times higher. Nevertheless, macroinvertebrate production was 3.7 times higher in Potamogeton stands. Greater production of epiphytic (50% higher) and benthic (84% higher) bacteria fulfilled additional food web inputs required in mesosaline wetlands, which had 50% higher dissolved organic carbon (DOC) and 84% higher sediment organic carbon available to bacteria. Dispersal limitation of amphipods after more frequent drawdowns in mesosaline wetlands may have promoted much higher biomass of competing chironomid larvae. Thus, vegetation type and direct algivory appeared less important to the structure and production of invertebrate food webs than were sediment organic matter, freshwater inflow that dilutes DOC as bacterial food, and the frequency of drawdowns that inhibit taxa with limited dispersal capability.
Keywords	Amphipoda ; Chara ; Chironomidae ; Potamogeton ; Stuckenia pectinata ; basins ; biomass ; dissolved organic carbon ; epiphytes ; flood irrigation ; food webs ; freshwater ; macroalgae ; macroinvertebrates ; microalgae ; sediments ; snowpack ; steppes ; vascular plants ; wetlands ; Wyoming
Language	English
Dates of publication	2023-03
Size	p. 457-472.
Publishing place	Springer US
Document type	Article ; Online
ZDB-ID	1428921-0
ISSN	1435-0629 ; 1432-9840
ISSN (online)	1435-0629
ISSN	1432-9840
DOI	10.1007/s10021-022-00768-3
Database	NAL-Catalogue (AGRICOLA)

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Article ; Online: Organic matter sources and flows in tundra wetland food webs.

Plesh, Steven P / Lovvorn, James R / Miller, Micah W C

PloS one

2023 Volume 18, Issue 5, Page(s) e0286368

Abstract: Arctic lowland tundra is often dominated by wetlands. As numbers and types of these wetlands change with climate warming, their invertebrate biomass and assemblages may also be affected. Increased influx of nutrients and dissolved organic matter (DOM) ... ...

Abstract	Arctic lowland tundra is often dominated by wetlands. As numbers and types of these wetlands change with climate warming, their invertebrate biomass and assemblages may also be affected. Increased influx of nutrients and dissolved organic matter (DOM) from thawing peat may alter the relative availability of organic matter (OM) sources, differentially affecting taxa with disparate dependence on those sources. In five shallow wetland types (<40 to 110 cm deep) and in littoral zones of deeper lakes (>150 cm), we used stable isotopes (δ13C, δ15N) to compare contributions of four OM sources (periphytic microalgae, cyanobacteria, macrophytes, peat) to the diets of nine macroinvertebrate taxa. Living macrophytes were not distinguishable isotopically from peat that likely contributed most DOM. Within invertebrate taxa, relative OM contributions were similar among all wetland types except deeper lakes. Physidae snails consumed substantial amounts of OM from cyanobacteria. However, for all other taxa examined, microalgae were the dominant or a major OM source (39-82%, mean 59%) in all wetland types except deeper lakes (20‒62%, mean 31%). Macrophytes and macrophyte-derived peat, likely consumed mostly indirectly as DOM-supported bacteria, ranged from 18‒61% (mean 41%) of ultimate OM sources in all wetland types except deeper lakes (38-80%, mean 69%). Invertebrate consumption of microalgal C may often have involved bacterial intermediates, or a mix of algae with bacteria consuming peat-derived OM. High production of periphyton with very low δ13C values were favored by continuous daylight illuminating shallow depths, high N and P levels, and high CO2 concentrations from bacterial respiration of peat-derived DOM. Although relative OM sources were similar across wetland types except deeper lakes, total invertebrate biomass was much higher in shallow wetlands with emergent vegetation. Impacts of warming on the availability of invertebrate prey to waterbirds will likely depend not on shifts in OM sources, but more on changes in overall number or area of shallow emergent wetlands.
MeSH term(s)	Animals ; Wetlands ; Food Chain ; Invertebrates ; Cyanobacteria ; Tundra ; Lakes ; Soil
Chemical Substances	Soil
Language	English
Publishing date	2023-05-26
Publishing country	United States
Document type	Journal Article ; Research Support, Non-U.S. Gov't
ZDB-ID	2267670-3
ISSN	1932-6203 ; 1932-6203
ISSN (online)	1932-6203
ISSN	1932-6203
DOI	10.1371/journal.pone.0286368
Database	MEDical Literature Analysis and Retrieval System OnLINE

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Article: Duck Use of Saline Wetlands Created by Irrigation in a Semiarid Landscape

Lovvorn, James R. / Crozier, Michelle L.

Wetlands. 2022 Jan., v. 42, no. 1

2022

Abstract: Throughout semiarid western North America, flood irrigation and associated small reservoirs have created or augmented many wetlands that otherwise would not exist or persist through summer. Diversion of mountain snowmelt from rivers has thereby created ... ...

Abstract	Throughout semiarid western North America, flood irrigation and associated small reservoirs have created or augmented many wetlands that otherwise would not exist or persist through summer. Diversion of mountain snowmelt from rivers has thereby created widely scattered hotspots of biodiversity. Increased urban water demands, higher profits from sprinkler irrigation, and climate-driven declines in mountain snowpack threaten these wetlands. Knowledge of unique functions of different wetland types and their spatial interactions would aid conservation of wetland complexes. We characterized use by ducks of wetlands with varying salinities, vegetation, nearby land use, and spatial relations in the Laramie Basin, Wyoming, USA. All duck species and social groups had higher densities in smaller wetlands. Pairs and broods of diving ducks and some dabbling ducks had highest densities in oligosaline wetlands (0.5–5 ‰ salinity) which have emergent plants for nesting cover. However, these ducks were commonly observed in mesosaline wetlands (5–18 ‰) which lack emergent cover but have higher availability of near-surface foods, suggesting differential use of wetland types for nesting and feeding. Accordingly, densities of some dabbling and diving ducks were higher when mesosaline wetlands were within 1 km. Hayfields or livestock grazing nearby seldom affected duck densities in wetlands, suggesting that with sparse upland cover in shortgrass steppe, many upland nesters sought cover in dry portions of the emergent fringe. For ducks in such intermountain basins, mesosaline wetlands with less stable water levels but high prey availability should be maintained in complexes near oligosaline wetlands with variably flooded emergent cover.
Keywords	basins ; biodiversity ; ducks ; flood irrigation ; highlands ; land use ; landscapes ; salinity ; snowmelt ; snowpack ; sprinkler irrigation ; steppes ; summer ; wetlands ; Wyoming
Language	English
Dates of publication	2022-01
Size	p. 4.
Publishing place	Springer Netherlands
Document type	Article
ZDB-ID	1097341-2
ISSN	1943-6246 ; 0277-5212
ISSN (online)	1943-6246
ISSN	0277-5212
DOI	10.1007/s13157-021-01525-3
Database	NAL-Catalogue (AGRICOLA)

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Article: Prey availability and foraging activity by tundra-nesting sea ducks: Strong preference for specific wetland types.

Miller, Micah W C / Lovvorn, James R / Graff, Nathan R / Stellrecht, Neesha C / Plesh, Steven P

Ecology and evolution

2023 Volume 13, Issue 9, Page(s) e10375

Abstract: Wetlands in Arctic tundra support abundant breeding waterbirds. Wetland types differing in area, depth, vegetation, and invertebrate biomass density may vary in importance to birds, and in vulnerability to climate change. We studied availability and use ... ...

Abstract	Wetlands in Arctic tundra support abundant breeding waterbirds. Wetland types differing in area, depth, vegetation, and invertebrate biomass density may vary in importance to birds, and in vulnerability to climate change. We studied availability and use of different wetland types by prelaying females of four species of sea ducks (Mergini) breeding on the Arctic Coastal Plain of Alaska, USA: long-tailed ducks (
Language	English
Publishing date	2023-09-20
Publishing country	England
Document type	Journal Article
ZDB-ID	2635675-2
ISSN	2045-7758
ISSN	2045-7758
DOI	10.1002/ece3.10375
Database	MEDical Literature Analysis and Retrieval System OnLINE

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Article: Feeding on epibenthic zooplankton by Long‐tailed Ducks: patch structure, profitability, and food web implications

Lovvorn, James R. / Brooks, Marjorie L.

Ecosphere. 2021 Nov., v. 12, no. 11

2021

Abstract: Models of the trophic needs and impacts of mobile predators in food web networks typically do not consider interactions between prey dispersion and threshold prey densities needed for profitable foraging. As a result, the biomass of predators that can be ...

Abstract	Models of the trophic needs and impacts of mobile predators in food web networks typically do not consider interactions between prey dispersion and threshold prey densities needed for profitable foraging. As a result, the biomass of predators that can be supported, or direct and indirect impacts of those predators on other food web components, may be poorly estimated. In Simpson Lagoon on the U.S. Arctic coast, epibenthic amphipods and especially mysid shrimp comprised most of the diets of fish, and of Long‐tailed Ducks (Clangula hyemalis) which strongly dominated prey consumption. Based on mean prey densities and gross estimates of prey biomass consumed, earlier studies estimated that continuous immigration of mysids into the lagoon was needed to support the predators. Comparison of variances from net samples vs. different probability density functions (PDFs) around reported means indicated that prey were not normally distributed, and instead occurred in patches with a skewed and wide range of densities. We used the functional response of captive Long‐tailed Ducks eating mysids, and a foraging energetics model, to identify minimum prey densities these diving ducks needed to feed profitably. Based on PDFs that assumed prey densities in all 1‐m² cells (patches for single feeding dives on potential mysid swarms) were independent, we simulated prey grids with slightly skewed to highly skewed distributions for the reported sample means. Results showed that the greater the range of patch densities, the smaller the fraction of prey biomass that was profitably available to the ducks, thereby increasing the requisite immigration of mysids. Consumption by ectothermic predators further decreased prey available and advanced the date and increased the amount of mysid influx required. The models also showed that the birds had to seek areas with non‐independent clusters of high‐density feeding patches as prey declined in late summer. Thus, spatially continuous measures of patch structure over larger scales are needed to estimate the fraction of prey that is functionally available. Our study emphasizes that when estimating trophic needs and impacts of mobile endothermic predators, simple means of prey density can be misleading, owing to interactions of prey patch structure with profitability thresholds.
Keywords	Amphipoda ; biomass ; coasts ; ectothermy ; endothermy ; fish ; food webs ; immigration ; models ; probability distribution ; profitability ; shrimp ; summer ; zooplankton ; Arctic region
Language	English
Dates of publication	2021-11
Publishing place	John Wiley & Sons, Ltd
Document type	Article
Note	JOURNAL ARTICLE
ZDB-ID	2572257-8
ISSN	2150-8925
ISSN	2150-8925
DOI	10.1002/ecs2.3780
Database	NAL-Catalogue (AGRICOLA)

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Article: Use of marine vs. freshwater proteins for egg‐laying and incubation by sea ducks breeding in Arctic tundra

Miller, Micah W. C. / Lovvorn, James R. / Graff, Nathan R. / Stellrecht, Neesha C.

Ecosphere. 2022 June, v. 13, no. 6

2022

Abstract: Understanding dietary nutrient sources is fundamental to conserving sensitive species, especially as climate change alters food web dynamics. Migratory species that depend on both marine and terrestrial habitats face unique challenges, as the locations ... ...

Abstract	Understanding dietary nutrient sources is fundamental to conserving sensitive species, especially as climate change alters food web dynamics. Migratory species that depend on both marine and terrestrial habitats face unique challenges, as the locations and quality of resources in the two realms may respond quite differently to environmental changes, with potential for spatial and temporal carryover effects. For sea ducks (Mergini) that winter at sea but move inland to breed, body size may determine their capacity to store nutrient reserves for later use in alternative habitats. We assessed ultimate sources of protein for reproduction in four sea duck species in northern Alaska: smaller‐bodied Long‐tailed Ducks and Steller's Eiders (Clangula hyemalis and Polysticta stelleri), and larger‐bodied Spectacled and King Eiders (Somateria fischeri and Somateria spectabilis). To assess the relative use of local freshwater foods vs. marine protein for both egg production and body maintenance of incubating females, we measured stable isotopes of carbon and nitrogen in egg membranes, red blood cells, marine and freshwater invertebrates, and vegetation. For egg production, isotope mixing models indicated that proteinaceous egg membranes of all four species were derived mostly (89%–95%) from freshwater foods on the breeding grounds, with broad individual variation in specific prey types selected by the larger species. For incubation, isotopes in red blood cells indicated that body maintenance of females also relied mainly (87%–91%) on freshwater foods in Long‐tailed Ducks and Steller's Eiders. However, incubating Spectacled and King Eiders obtained only about 60% of their protein from freshwater foods and the remainder from marine‐derived body tissues. The latter strategy allows the larger‐bodied species to incubate almost continuously, whereas the smaller species must take more frequent incubation breaks and generally incur higher rates of predation on eggs. Thus, depending on body size, cross‐seasonal effects of feeding conditions in marine habitats may strongly influence population processes well after the birds move to inland nesting sites. Although conservation programs on land and sea are often researched, planned, and administered by different agencies and organizations, our results emphasize the need to coordinate marine and land‐based efforts for species that integrate conditions across both environments.
Keywords	Somateria spectabilis ; body size ; carbon ; climate change ; ducks ; egg production ; eggs ; food composition ; food webs ; freshwater ; migratory species ; nitrogen ; oviposition ; predation ; tundra ; winter ; Alaska
Language	English
Dates of publication	2022-06
Publishing place	John Wiley & Sons, Inc.
Document type	Article
Note	JOURNAL ARTICLE
ZDB-ID	2572257-8
ISSN	2150-8925
ISSN	2150-8925
DOI	10.1002/ecs2.4138
Database	NAL-Catalogue (AGRICOLA)

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Article: Detecting silent stressors: Trace element effects on nutritional status of declining scoter ducks of Puget Sound, USA

Brooks, Marjorie L / Lovvorn, James R / Behnke, Jessica Hallman / Anderson, Eric M

Science of the total environment. 2021 Apr. 20, v. 766

2021

Abstract: White-winged scoters (Melanitta fusca; WWSC) and surf scoters (M. perspicillata; SUSC) have declined by over 60% in recent decades. Identifying contributing factors from within a mosaic of sublethal, multiple stressors is challenging. In urbanized Puget ... ...

Abstract	White-winged scoters (Melanitta fusca; WWSC) and surf scoters (M. perspicillata; SUSC) have declined by over 60% in recent decades. Identifying contributing factors from within a mosaic of sublethal, multiple stressors is challenging. In urbanized Puget Sound, Washington, USA where scoters winter, changes in prey availability explained only a portion of local declines, suggesting that other “silent stressors” such as sublethal contaminants might play a role. Past studies of pollutant effects on scoters used Fisherian statistics that often revealed few correlates; however, novel statistical approaches could detect and provide more insights about sublethal impacts. Our objectives were to (1) relate pollutant accumulation to health of the birds, and (2) compare permutational multivariate statistics with traditional approaches in identifying sublethal health effects. We collected scoters from three locations in Puget Sound in December 2005 and March 2006, and measured cadmium (Cd), mercury (Hg), and selenium (Se) levels in livers and kidneys. To assess impacts of low contaminants levels in tissues on nutritional status (whole-body mass, lipid, and protein; and triglycerides, β-hydroxybutyrate, and uric acid in blood), we compared statistical methods. Permutational multivariate methods use Monte Carlo techniques to assess how an integrated matrix of physiological responses in each animal respond to contaminants. Univariate regressions revealed very few and inconsistent relationships. In contrast, multivariate models showed that liver Hg and Se explained 25% of the variance in nutritional status of white-winged scoters; and in surf scoters, Cd, Hg, and Se in tissues explained 14 to 27% of nutritional status depending on site. The influence of these factors equals other aspects of habitat such as foraging conditions. Our study indicates that permutational multivariate statistics can be a powerful tool for identifying sublethal contaminant associations that, with non-contaminant stressors, can influence nutritional status and thus, contribute to population dynamics.
Keywords	blood ; cadmium ; environment ; habitats ; liver ; mercury ; multivariate analysis ; nutritional status ; pollutants ; population dynamics ; selenium ; urbanization ; uric acid ; variance ; winter ; Puget Sound
Language	English
Dates of publication	2021-0420
Publishing place	Elsevier B.V.
Document type	Article
Note	NAL-AP-2-clean
ZDB-ID	121506-1
ISSN	1879-1026 ; 0048-9697
ISSN (online)	1879-1026
ISSN	0048-9697
DOI	10.1016/j.scitotenv.2020.144247
Database	NAL-Catalogue (AGRICOLA)

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Article ; Online: Detecting silent stressors: Trace element effects on nutritional status of declining scoter ducks of Puget Sound, USA.

Brooks, Marjorie L / Lovvorn, James R / Behnke, Jessica Hallman / Anderson, Eric M

The Science of the total environment

2020 Volume 766, Page(s) 144247

Abstract	White-winged scoters (Melanitta fusca; WWSC) and surf scoters (M. perspicillata; SUSC) have declined by over 60% in recent decades. Identifying contributing factors from within a mosaic of sublethal, multiple stressors is challenging. In urbanized Puget Sound, Washington, USA where scoters winter, changes in prey availability explained only a portion of local declines, suggesting that other "silent stressors" such as sublethal contaminants might play a role. Past studies of pollutant effects on scoters used Fisherian statistics that often revealed few correlates; however, novel statistical approaches could detect and provide more insights about sublethal impacts. Our objectives were to (1) relate pollutant accumulation to health of the birds, and (2) compare permutational multivariate statistics with traditional approaches in identifying sublethal health effects. We collected scoters from three locations in Puget Sound in December 2005 and March 2006, and measured cadmium (Cd), mercury (Hg), and selenium (Se) levels in livers and kidneys. To assess impacts of low contaminants levels in tissues on nutritional status (whole-body mass, lipid, and protein; and triglycerides, β-hydroxybutyrate, and uric acid in blood), we compared statistical methods. Permutational multivariate methods use Monte Carlo techniques to assess how an integrated matrix of physiological responses in each animal respond to contaminants. Univariate regressions revealed very few and inconsistent relationships. In contrast, multivariate models showed that liver Hg and Se explained 25% of the variance in nutritional status of white-winged scoters; and in surf scoters, Cd, Hg, and Se in tissues explained 14 to 27% of nutritional status depending on site. The influence of these factors equals other aspects of habitat such as foraging conditions. Our study indicates that permutational multivariate statistics can be a powerful tool for identifying sublethal contaminant associations that, with non-contaminant stressors, can influence nutritional status and thus, contribute to population dynamics.
MeSH term(s)	Animals ; Ducks ; Environmental Pollutants/analysis ; Environmental Pollutants/toxicity ; Nutritional Status ; Selenium/analysis ; Selenium/toxicity ; Trace Elements ; Washington
Chemical Substances	Environmental Pollutants ; Trace Elements ; Selenium (H6241UJ22B)
Language	English
Publishing date	2020-12-24
Publishing country	Netherlands
Document type	Journal Article
ZDB-ID	121506-1
ISSN	1879-1026 ; 0048-9697
ISSN (online)	1879-1026
ISSN	0048-9697
DOI	10.1016/j.scitotenv.2020.144247
Database	MEDical Literature Analysis and Retrieval System OnLINE

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Article: Sustaining ecological and subsistence functions in conservation areas: eider habitat and access by Native hunters along landfast ice

LOVVORN, JAMES R / JEWETT, STEPHEN C / MAHONEY, ANDREW H / ROCHA, AARIEL R

Environmental conservation. 2018 Dec., v. 45, no. 4

2018

Abstract: In the Arctic, rapid climate change has kindled efforts to delineate and project the future of important habitats for marine birds and mammals. These animals are vital to subsistence economies and cultures, so including the needs of both animals and ... ...

Abstract	In the Arctic, rapid climate change has kindled efforts to delineate and project the future of important habitats for marine birds and mammals. These animals are vital to subsistence economies and cultures, so including the needs of both animals and hunters in conservation planning is key to sustaining social-ecological systems. In the northeast Chukchi Sea, a nearshore corridor of open water is a major spring migration route for half a million eider ducks that are hunted along the landfast ice. Zoning areas for industrial activities or conservation should consider both eider habitat and hunter access to those habitats from the variable ice edge. Based on benthic sampling in 2010‒2012, a model of eider foraging energetics and satellite data on ice patterns in April and May 1997‒2011, we mapped the range of positions of the landfast ice edge relative to a given dispersion of habitat suitable for eider feeding. In some sectors, feeding areas were too limited or too far from landfast ice to provide regular hunting access. In other sectors, overlap of the ice edge with eider feeding habitat was quite variable, but often within a consistent geographic range. Areas accessible to hunters were a small fraction of total eider habitat, so areas adequate for conserving eiders would not necessarily include areas that meet the hunters’ needs. These results can inform spatial planning of industrial activities that yield cash income critical to subsistence hunting in less developed locations. Our study provides an approach for mapping ‘subsistence conservation areas’ throughout the Arctic and an example for such efforts elsewhere.
Keywords	climate change ; conservation areas ; ducks ; foraging ; geographical distribution ; habitats ; hunters ; ice ; income ; mammals ; models ; planning ; remote sensing ; spring ; water birds ; zoning ; Arctic region ; Chukchi Sea
Language	English
Dates of publication	2018-12
Size	p. 361-369.
Publishing place	Cambridge University Press
Document type	Article
ZDB-ID	1470226-5
ISSN	1469-4387 ; 0376-8929
ISSN (online)	1469-4387
ISSN	0376-8929
DOI	10.1017/S0376892918000103
Database	NAL-Catalogue (AGRICOLA)

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Article ; Online: Effects of air and water temperatures on resting metabolism of auklets and other diving birds.

Richman, Samantha E / Lovvorn, James R

Physiological and biochemical zoology : PBZ

2011 Volume 84, Issue 3, Page(s) 316–332

Abstract: For small aquatic endotherms, heat loss while floating on water can be a dominant energy cost, and requires accurate estimation in energetics models for different species. We measured resting metabolic rate (RMR) in air and on water for a small diving ... ...

Abstract	For small aquatic endotherms, heat loss while floating on water can be a dominant energy cost, and requires accurate estimation in energetics models for different species. We measured resting metabolic rate (RMR) in air and on water for a small diving bird, the Cassin's auklet (Ptychoramphus aleuticus), and compared these results to published data for other diving birds of diverse taxa and sizes. For 8 Cassin's auklets (~165 g), the lower critical temperature was higher on water (21 °C) than in air (16 °C). Lowest values of RMR (W kg⁻¹) averaged 19% higher on water (12.14 ± 3.14 SD) than in air (10.22 ± 1.43). At lower temperatures, RMR averaged 25% higher on water than in air, increasing with similar slope. RMR was higher on water than in air for alcids, cormorants, and small penguins but not for diving ducks, which appear exceptionally resistant to heat loss in water. Changes in RMR (W) with body mass either in air or on water were mostly linear over the 5- to 20-fold body mass ranges of alcids, diving ducks, and penguins, while cormorants showed no relationship of RMR with mass. The often large energetic effects of time spent floating on water can differ substantially among major taxa of diving birds, so that relevant estimates are critical to understanding their patterns of daily energy use.
MeSH term(s)	Air ; Animals ; Basal Metabolism ; Birds/physiology ; Body Size ; Body Temperature Regulation ; Charadriiformes/physiology ; Ducks/physiology ; Spheniscidae/physiology ; Temperature ; Water
Chemical Substances	Water (059QF0KO0R)
Language	English
Publishing date	2011-05
Publishing country	United States
Document type	Comparative Study ; Journal Article ; Research Support, Non-U.S. Gov't
ZDB-ID	1473845-4
ISSN	1537-5293 ; 1522-2152
ISSN (online)	1537-5293
ISSN	1522-2152
DOI	10.1086/660008
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