Development of water level regulation strategies for fish and wildlife,upper Mississippi river system

Water level regulation has been proposed as a tool for maintaining or enhancing fish and wildlife resources in navigation pools and associated flood plains of the Upper Mississippi River System. Research related to the development of water level management plans is being conducted under the Long Term Resource Monitoring Program. Research strategies include investigations of cause and effect relationships, spatial and temporal patterns of resource components, and alternative problem solutions. The principal hypothesis being tested states that water level fluctuations resulting from navigation dam operation create less than optimal conditions for the reproduction and growth of target aquatic macrophyte and fish species. Representative navigation pools have been selected to describe hydrologic, engineering, and legal constraints within which fish and wildlife objectives can be established. Spatial analyses are underway to predict the magnitude and location of habitat changes that will result from controlled changes in water elevation.     

Primary production and light dynamics in an upper Mississippi River backwater

In nutrient-rich systems, phytoplankton production is frequently light-limited and light attenuation can become a critical factor controlling the rates of production, especially in shallow systems subject to wind resuspension of sediments. This study examined the relation between light availability and primary production in Lake Onalaska, a shallow, nutrient-rich impoundment of the Upper Mississippi River. Continuous water quality monitors recorded dissolved oxygen, temperature and light flux at a network of sites from July to September 1990 and these data were used to estimate the primary production, light availability and light attenuation coefficients. The gross primary production averaged 2·2gCm^?2 day^?1 and was due almost entirely to phytoplankton. Production was strongly light-limited, with most of the light attenuation due to non-algal components, possibly related to wind resuspension in this shallow system. Regression analyses revealed significant effects of wind speed and direction on light attenuation.     

Analysis of water level management on the upper Mississippi River (1980–1990)

Management at 27 low-head dams affects water surface elevations for a 1050km stretch of the Upper Mississippi River (UMR) between St Louis, Missouri and Minneapolis, Minnesota. A systemic overview is given of current operating plans at dams on the UMR and historical data are analysed to determine how well the operating plans are being met. Water level elevations at all 27 dams are regulated as a function of discharge, although plans are specific for each dam. The management objective is to maintain a target water level at specific locations (control point) in each impoundment over specific ranges of discharge. The target water level and control point may change as discharge changes in each impoundment. In some of the impoundments water regulation causes drawdowns below the elevation for which the dams were planned, and at other dams no drawdown occurs. During the navigation seasons of 1980 to 1990, water levels were within their target window for an average of 72·5% of the time for 25 dams analysed. Difficulties in meeting targets are caused by winds, local rainfall events, ice dams and rapidly fluctuating discharges from tributaries with upstream reservoirs used for peaking hydropower.     

Habitat rehabilitation on the upper Mississippi River

Upper Mississippi River ecological integrity has been severely compromised by human activity during the last 50 years. In response to the continuing decline of natural resource values, two approaches for protecting and improving the Upper Mississippi River-floodplain ecosystem have been used. Habitat rehabilitation and enhancement projects are being constructed at 54 locations to provide site-specific rehabilitation. The projects are designed to counteract the adverse ecological effects of sedimentation through (1) flow introductions, (2) the isolation of backwaters; and (3) flow diversions and wave breaks. Channel maintenance projects are being re-evaluated in an attempt to construct or modify existing river training structures that are environmentally sympathetic. The latter approach works with the river's energy, whereas the former attempts to overcome riverine processes. Both approaches have significant limitations because they affect limited areas. A proposal is presented that restores some ecosystem integrity by re-establishing occasional low river stages that occurred before the implementation of the Upper Mississippi River Navigation System.     

Spatial patterns of fish communities in the Upper Mississippi River System: assessing fragmentation by low‐head dams

We assessed the similarity of fish communities among river reaches to assess community‐level fragmentation by low‐head dams in the Upper Mississippi River System (UMRS). The spatial coverage of standardized electrofishing sampling used in the Long Term Resource Monitoring Program (LTRMP) was extended for three of the six regional trend areas (RTA; pools 4, 13, and the Open River Reach) to include river reaches (outpools) immediately upstream and downstream from the standard RTA from 15 June to 31 October 2000. Additionally, pools 19 and 20 were sampled in September 2000. Cluster analysis and non‐metric multidimensional scaling of community composition and structure data revealed two major groups, upper and lower reaches, and four (for composition) or five (for structure) sub‐groupings of river reaches. In general, all outpools grouped with the nearest RTA for both community composition (no exception) and community structure (one exception). This suggests that fragmentation of fish communities from low‐head dams is minimal. Mantel correlations demonstrated strong inverse association between the similarity of fish communities with the distance between reaches. Habitat variables measured during electrofishing collections were significantly correlated with spatial variation of fish composition and community structure, but provided only marginal improvements to correlations with distance between reaches alone. Furthermore, habitat variability among river reaches also was related to distance between reaches. Determining the extent to which variation of fish communities is related to habitat or demographic processes (e.g. migration, larval drift, source‐sink dynamics) will be challenging for this system. Although low‐head dams on the UMRS may restrict movements for individuals and populations of certain fish species, we found little evidence that these effects have led to substantial, community‐level fragmentation. Copyright © 2006 John Wiley & Sons, Ltd.     

Light availability and growth of wildcelery (Vallisneria americana) in upper Mississippi River backwaters

Large beds of Vallisneria americana declined in the backwaters of the Upper Mississippi River after a drought that occurred between 1987 and 1989. One hypothesis for this decline is that low light availability may have decreased net photosynthesis to the extent that overwintering tubers were not formed. Following the decline, light availability remained low. To determine what light levels would be necessary for the re-establishment of Vallisneria in the Upper Mississippi River, the long-term growth of plants in a backwater lake and in an experimental pond was measured while the surface and subsurface light were monitored continuously. Plants grown from tubers transplanted to 0·5, 1·0 and l·5m depth in the lake grew and produced tubers only at 0·5m depth (9% of surface light). At 1·0 m, light availability was less than 1% of the surface light. Plants grown from tubers in experimental ponds with four shade treatments (2, 5, 9 and 25% of surface light) for the same growing period produced replacement-weight tubers in 9% light. For a longer growing season, plants also produced replacement-weight tubers in treatments with at least 5% of surface light. An average light-extinction coefficient of 4·64 m^?1 was calculated for the backwater lake based on continuous data collected during 94 days during the growing season from eight widely separated sites. Using equations based on the average extinction coefficient for the lake and average leaf lengths of plants grown in experimental ponds, we predict that in years with comparable turbidity, plants grown from locally collected tubers will grow and produce replacement tubers only at depths of 0·8 m or less.     

Declines in aquatic vegetation in navigation pool no. 8, upper Mississippi River between 1975 and 1991

The biomass and frequency of occurrence of aquatic macrophytes observed during 1975 and 1991 in Navigation Pool No. 8 of the upper Mississippi River were compared. Samples were collected from contiguous backwater, isolated backwater and impounded habitat types. Overall declines were observed in both frequency and biomass between the two years. Changes in depth, water clarity and bed sediment texture were also observed. About 50% of the samples collected in 1991 had no vegetation, compared with 20% in 1975. The greatest reduction in frequency occurred among submergent taxa (77% in 1975 to 31% in 1991), whereas only slight declines were observed for emergent taxa. However, the mean biomass of both lifeforms decreased substantially (48 to 1 g/m² for submergent and 61 to 27 g/m² for emergent lifeforms). The mean total biomass declined from 114g/m² in 1975 to 35 g/m² in 1991. Declines were most evident in the impounded habitat, where the frequency of occurrence of submergent taxa decreased from 83% in 1975 to 11% in 1991, and mean biomass decreased from 90 to 1 g/m². Similar but less extensive reductions in frequency and biomass of submergent taxa were observed in the contiguous backwater habitat. In contrast, both the frequency and biomass of floating-leaved macrophytes increased slightly in contiguous backwater habitats. Changes in frequency and biomass of macrophytes in isolated backwater habitats were neligible between 1975 and 1991. It is hypothesized that changes in the macrophyte community structure may be related to physical changes associated with 57 years of impoundment.     

Nitrogen dynamics in sediment during water level manipulation on the Upper Mississippi River

Nitrogen (N) has been linked to increasing eutrophication in the Gulf of Mexico and as a result there is increased interest in managing and improving water quality in the Mississippi River system. Water level reductions, or ‘drawdowns’, are being used more frequently in large river impoundments to improve vegetation growth and sediment compaction. We selected two areas of the Upper Mississippi River system (Navigation Pool 8 and Swan Lake) to examine the effects of water level drawdown on N dynamics. Navigation Pool 8 experienced summer drawdowns in 2001 and 2002. Certain areas of Swan Lake have been drawn down annually since the early 1970s where as other areas have remained inundated. In the 2002 Pool 8 study we determined the effects of sediment drying and rewetting resulting from water level drawdown on (1) patterns of sediment nitrification and denitrification and (2) concentrations of sediment and surface water total N (TN), nitrate, and ammonium (NH). In 2001, we only examined sediment NH and TN. In the Swan Lake study, we determined the long‐term effects of water level drawdowns on concentrations of sediment NH and TN in sediments that dried annually and those that remained inundated. Sediment NH decreased significantly in the Pool 8 studies during periods of desiccation, although there were no consistent trends in nitrification and denitrification or a reduction in total sediment N. Ammonium in sediments that have dried annually in Swan Lake appeared lower but was not significantly different from sediments that remain wet. The reduction in sediment NH in parts of Pool 8 was likely a result of increased plant growth and N assimilation, which is then redeposited back to the sediment surface upon plant senescence. Similarly, the Swan Lake study suggested that drawdowns do not result in long term reduction in sediment N. Water level drawdowns may actually reduce water retention time and river‐floodplain connectivity, while promoting significant accumulation of organic N. These results indicate that water level drawdowns are probably not an effective means of removing N from the Upper Mississippi River system. Copyright © 2006 John Wiley & Sons, Ltd.     

Shallow and deep water aquatic vegetation potential for a midlatitude pool of the Upper Mississippi River System with drawdown

Prior to navigation dam and levee placement, the Upper Mississippi River (UMR) flowed through a wide floodplain supporting a diverse ecosystem. Diversity was created by variable flood frequencies and water flow, but presently high and static water levels supporting river navigation have caused low diversity of aquatic vegetation in locations within the UMR. A pool‐scale water level drawdown was proposed as a wetland management tool to mimic historic low water flow for UMR Navigation Pool 18, between Oquawka and Keithsburg, IL. The objectives of this research are to determine plant species, density, and diversity expected for a drawdown in Pool 18. A seedbank and propagule assay was used to evaluate drawdown plant species response. Emergence was tested using river bottom substrate samples collected in 2009 from the proposed drawdown area. Samples were treated at two hydrologic levels: shallow (3‐cm depth) and deep (16‐cm depth). Dominant species in the shallow flooded treatment were Gratiola neglecta, Leersia oryzoides, Eleocharis palustris, Sagittaria latifolia, and Ammania coccinea. Deep flooded dominant taxa included G. neglecta, S. latifolia, Vallisneria americana, and A. coccinea. Each treatment indicated a seedbank of moderate diversity with a shallow treatment diversity of D = 0.56 and deep treatment diversity of D = 0.44. Plant density for the shallow flooded treatment was 213 stems/m² (±112; 95% CI), and deep flooded hydrologic treatment, 206 stems/m² (±82; 95% CI). It is expected that this drawdown will provide an intermediate ecological disturbance resulting in greater species diversity and density currently lacking in this portion of the Upper Mississippi River System.     

Contaminant concentrations in bald eagles nesting on Lake Superior,the upper Mississippi River,and the St. Croix River

We measured concentrations of DDE, total PCBs, and mercury in bald eagle (Haliaeetus leucocephalus) nestlings at three locations in the upper Midwest: Lake Superior, the upper Mississippi River, and the St. Croix River, 2006–2008. We also analyzed trends in concentrations of these contaminants for eagles on the southern shore of Lake Superior, from 1989 to 2008, using the current and previously published data. Concentrations of DDE in nestling blood plasma samples were greatest on Lake Superior (geometric mean: 16.2 μg/kg, n = 29), whereas concentrations of total PCBs were highest in Mississippi River samples (88.6 μg/kg, n = 51). Mercury concentrations were highest along the upper St. Croix River (6.81 μg/g wet weight in feathers, n = 19). For Lake Superior, DDE concentrations declined significantly in nestling blood plasma samples from 1989 to 2008, an average of 3.0% annually. Similarly, total PCBs in Lake Superior eaglets decreased 4.0% annually from 1989 to 2008, and mercury concentrations in nestling feathers from Lake Superior nests also decreased significantly from 1991 to 2008, 2.4% per year. With the possible exception of mercury on the upper St. Croix River, mean concentrations in 2006–2008 of all three compounds were below levels associated with significant impairment of reproduction for all sites, and reproductive rates at all three sites averaged > 1.2 young per occupied territory, which is greater than the rate indicative of a healthy population.     

滦河流域大中型闸坝水文生态效应

为揭示滦河流域水库对下游河流水文影响,辨析其生态效应,建立了流域闸坝水生态效应评估体系,运用河流影响因子(RI)法评估了闪电河、庙宫、潘家口和桃林口水库的水生态效应,进一步运用水文变化范围(RVA)法评估了潘家口水库的水生态效应。结果表明:1各水库对水文的影响程度为:潘家口桃林口闪电河庙宫,水库的水文效应同时受其级别(库容)和河流原始径流量影响,小型河流水库的水文生态效应不容忽视。2潘家口水库IHA指标(RVA法)总改变度为0.88,第1~2组指标发生了高度改变(0.91),第3~5组指标发生中度改变(0.45),强烈改变了鱼类洄游、底栖生物和植物群落等生态过程。生态水文同步监测与生态模型构建将有助于提升河流生态系统预警能力,为河流生态恢复提供科学依据和技术工具。     

Spatial and Temporal Dynamics of Suspended Particle Characteristics and Composition in Navigation Pool 19 of the Upper Mississippi River

Suspended particles are an essential component of large rivers influencing channel geomorphology, biogeochemical cycling of nutrients, and food web resources. The Upper Mississippi River is a large floodplain river that exhibits pronounced spatiotemporal variation in environmental conditions and biota, providing an ideal environment for investigating dynamics of suspended particles in large river ecosystems. Here we investigated two questions: (i) How do suspended particle characteristics (e.g. size and morphology) vary temporally and spatially? and (ii) What environmental variables have the strongest association with particle characteristics? Water sampling was conducted in June, August, and September of 2013 and 2014 in Navigation Pool 19 of the Upper Mississippi River. A FlowCAM® (Flow Cytometer and Microscope) particle imaging system was used to enumerate and measure particles 53–300 μm in diameter for size and shape characteristics (e.g. volume, elongation, and symmetry). Suspended particle characteristics varied considerably over space and time and were strongly associated with discharge and concentrations of nitrate + nitrite (NO₃^?) and soluble reactive phosphorus. Particle characteristics in backwaters were distinct from those in other habitats for most of the study period, likely due to reduced hydrologic connectivity and higher biotic production in backwaters. During low discharge, phytoplankton and zooplankton made up relatively greater proportions of the observed particles. Concurrently during low discharge, concentrations of chlorophyll, volatile suspended solids, and total phosphorus were higher. Our results suggest that there are complex interactions among space, time, discharge, and other environmental variables (e.g. water nutrients), which drive suspended particle dynamics in large rivers. Copyright © 2017 John Wiley & Sons, Ltd.     

Responses of consumers and food resources to a high magnitude,unpredicted flood in the upper Mississippi River basin

The Mississippi and Missouri Rivers experienced flooding in 1993 that fell outside the annual predictable flood period of spring and early summer. Flooding began in late June, peaked in late July (25 232 m³/s on the upper Mississippi and 21 240 m³/s on the Missouri) and remained at or near flood stage into October 1993. This study was performed to determine if disturbance by an unpredicted flood event would alter trophic dynamics of river–floodplain systems by creating shifts in the composition of organic matter available to consumers. The Ohio River, which did not flood during the same period, was examined for comparison. Stable isotopic ratios of carbon and nitrogen from samples collected in 1993 and 1994 were used to characterize potential food sources and determine linkages between food sources and invertebrate and fish consumers. Pairwise contrasts, performed separately for each river, indicated there were few interannual differences in δ¹³C and δ¹⁵N of organic matter sources and consumers. Between sample period (flood year versus normal water year) trends in both flooded rivers were similar to between‐year trends observed for the Ohio River. Trophic structure of the Mississippi and Ohio Rivers was similar in both years, with fine and ultra‐fine transported organic matter and dissolved organic matter representing the major sources of organic matter. Overlapping isotopic signatures in the Missouri River made tracking of sources through the consumers difficult, but similarities in δ¹³C and δ¹⁵N between years indicated trophic structure did not change in response to the flood. The results suggest that consumers continued to rely on sources of organic matter that would be used in the absence of the unpredicted 1993 flood. It is proposed that trophic structure did not change in response to flooding in the Mississippi and Missouri Rivers because both rivers exhibited the same trends observed in the Ohio River. Copyright © 2001 John Wiley & Sons, Ltd.     

Influence of a high‐head dam as a dispersal barrier to fish community structure of the Upper Mississippi River

In river systems, high‐head dams may increase the distance‐decay of fish community similarity by creating nearly impermeable dispersal barriers to certain species from upstream reaches. Substantial evidence suggests that migratory species are impacted by dams, and most previous studies in stream/river networks have focused on small streams and headwaters. Here, we assess whether a high‐head dam (Lock and Dam 19; LD 19) on a large river, the Upper Mississippi River (UMR), substantially alters fish community structure relative to variability expected to occur independent of the dam's effect as a fish dispersal barrier. Using fish catch per unit effort data, we modelled the distance‐decay function for the UMR fish community and then estimated the similarity that would be expected to occur across LD19 and compared it with measured similarity. Measured similarity in the fish community above and below LD19 was close to the expected value based on the distance‐decay function, suggesting LD19 does not create an abrupt transition in the fish community. Although some migratory fish species no longer occur above LD19 (e.g., skipjack herring, Alosa chrysochloris), these species do not occur in high abundance below the dam and so do not drive variation in fish community structure. Instead, much of the variation in species structure is driven by the loss/gain of species across the latitudinal gradient. Lock and Dam 19 does not appear to be a clear transition point in the river's fish community, although it may function as a meaningful barrier for particular species (e.g., invasive species) and warrant future attention from a management perspective.     

Morphological changes of the Lower Mississippi River: geomorphological response to engineering intervention

During the twentieth century, the planform and profile of the Lower Mississippi River from Cairo, Illinois, to New Orleans, Louisiana, have been transformed by a series of engineering modifications. These include steepening of the long profile by removal of the most sinuous bends, extensive bank stabilization, and regulating sediment movement by dyke field construction. Prior to these modifications, the Lower Mississippi River adjusted its morphology in the planform, long profile and cross‐section. Planform adjustment has, however, effectively been negated and other adjustments are now constrained. Nevertheless, analysis of hydrographic surveys between 1949 and 1989 demonstrates that geomorphological response during the post‐cutoff period remained complex. Morphological adjustments involved phased patterns of aggradation and degradation, together with changes in cross‐sectional form, and in the number, size, location and shape of pools and crossings. Greatest changes occurred in the early post‐cutoff period (1949–64) upstream from Vicksburg, Mississippi, but were accompanied by complementary changes elsewhere which propagated downstream. The combined set of responses may be interpreted with respect to a dynamic equilibrium in which the river responded to additional energy created by the cutoffs by increasing and adapting flow resistance over various scales and time periods. This study helps resolve paradoxes from previous analyses, and has significance for interpreting past engineering impacts and for suggesting future management strategies for the Lower Mississippi River. Copyright © 2005 John Wiley & Sons, Ltd.     

Reproductive success of the interior least tern (Sterna antillarum) in relation to hydrology on the Lower Mississippi River

The annual hydrograph of large rivers, including flood pulses and low‐flow periods, is believed to play a primary role in the productivity of biota associated with these ecosystems. We investigated the relationship between river hydrology and Interior least tern (Sterna antillarum) reproductive success on the Lower Mississippi River from April to July 1986–1993. The number of fledglings produced per adult pair was negatively correlated with July mean (r=?0.95, p=0.0004) and July maximum river elevation (r=?0.97, p=0.0001), but no other aspects of river hydrology were related to tern reproduction. Low‐water elevations in July may benefit least tern reproductive success by increasing sand island area or the area of shallow‐water habitat that surrounds islands. Loss of deep‐water habitats in conjunction with an increase in shallow habitat during stage decreases may concentrate fish prey in shallow‐water habitats and backwater areas, thereby increasing food availability during chick‐rearing. Copyright © 2002 John Wiley & Sons, Ltd.     

The Historical Struggle with Floods on the Mississippi River Basin

For the past 150 years a titanic struggle has been underway between the people of the Mississippi River basin and its floods. Humans first tried to control the floods with structures, channel straightening, levees, dams, and reservoirs, but after 100 years and the expenditure of billions of dollars; losses to property and lives continued to grow. Efforts since the 1950s to encourage land-use changes in flood-prone areas and the use of flood insurance often have been thwarted by government relief and continuing human desire to reside in floodplains. During the massive floods in 1993, 1996, and 1997 only ten percent of the residents of flooded areas had flood insurance. The river system brings enormous economic value to the United States, and interests in protecting and enhancing the natural environment of the river are often in direct conflict with economic interests. Flood mitigation is often caught in the middle of the debate about how to manage the river system to satisfy all interests. Recent floods have taught several key lessons about environmental blessings and dangers, and reveal that the nation's policies regarding flood mitigation/relief must change with more emphasis on personal responsibility. The huge and costly infrastructure built to control the major rivers for floods and navigation is aging and will soon need replacement, offering an opportunity to better handle future flood mitigation to satisfy the basin's complex mix of economic, human, and environmental interests.     

Impacts of the Mississippi River spillway opening on faecal coliform concentration in Lake Pontchartrain

This study focuses on explaining the role of the periodic opening of the Bonnet Carré Spillway on the faecal coliform levels in Lake Pontchartrain. Water quality data were collected by the Lake Pontchartrain Basin Foundation to analyse the faecal coliform bacteria levels, turbidity, salinity, temperature, and dissolved oxygen in the lake water. The results show that the faecal coliform concentrations were lower during the spillway opening than the preceding and subsequent months. Statistical analysis shows the dilution effect, characterized by an effective reduction in faecal coliform concentrations due to increase in water turbidity, coupled with decrease in water salinity and dissolved oxygen because of spillway opening. It is also indicated that the combine effect of more than one factor produces a significantly greater reduction in the faecal coliform than each factor acting independently. Periodic spikes in faecal coliform concentrations were observed at sampling points located near the outlets of streams and drainage canals with high settlement densities. A positive correlation (r = 0.78467) was found between the average faecal coliform counts and average monthly precipitations, highlighting the role of precipitation run‐offs. The results of this study indicate that the spillway opening contributed to a reduction in the faecal coliform concentration in the lake's surface water during the studied periods.     

Spatial and Temporal Relationships Between the Invasive Snail Bithynia tentaculata and Submersed Aquatic Vegetation in Pool 8 of the Upper Mississippi River

Bithynia tentaculata is an invasive snail that was first reported in Lake Michigan in 1871 and has since spread throughout a number of freshwater systems of the USA. This invasion has been extremely problematic in the Upper Mississippi River as the snails serve as intermediate hosts for several trematode parasites that have been associated with waterfowl mortality in the region. This study was designed to assess the abundance and distribution of B. tentaculata relative to submersed aquatic vegetation as macrophytes provide important nesting and food resources for migrating waterfowl. Temporal changes in both vegetation and snail densities were compared between 2007 and 2015. Between these years, B. tentaculata densities have nearly quadrupled despite minor changes in vegetation abundance, distribution and composition. Understanding the spatial distribution of B. tentaculata in relation to other habitat features, including submersed vegetation, and quantifying any further changes in the abundance and distribution of B. tentaculata over time will be important for better identifying areas of risk for disease transmission to waterfowl. Copyright © 2017 John Wiley & Sons, Ltd.     

Projected Risk of Population Declines for Native Fish Species in the Upper Mississippi River

Conservationists are in need of objective metrics for prioritizing the management of habitats. For individual species, the threat of extinction is often used to prioritize what species are in need of conservation action. Using long‐term monitoring data, we applied a Bayesian diffusion approximation to estimate quasi‐extinction risk for 54 native fish species within six commercial navigation reaches along a 1350‐km gradient of the upper Mississippi River system. We found a strong negative linear relationship between quasi‐extinction risk and distance upstream. For some species, quasi‐extinction estimates ranged from nearly zero in some reaches to one in others, suggesting substantial variability in threats facing individual river reaches. We found no evidence that species traits affected quasi‐extinction risk across the entire system. Our results indicate that fishes within the upper Mississippi River system face localized threats that vary across river impact gradients. This suggests that conservation actions should be focused on local habitat scales but should also consider the additive effects on downstream conditions. We also emphasize the need for identification of proximate mechanisms behind observed and predicted population declines, as conservation actions will require mitigation of such mechanisms. Published 2014. This article is a U.S. Government work and is in the public domain in the USA.