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Denitrification across landscapes and waterscapes: A synthesis

Denitrification is a critical process regulating the removal of bioavailable nitrogen (N) from natural and human-altered systems. While it has been extensively studied in terrestrial, freshwater, and marine systems, there has been limited communication among denitrification scientists working in these individual systems. Here, we compare rates of denitrification and controlling factors across a range of ecosystem types. We suggest that terrestrial, freshwater, and marine systems in which denitrification occurs can be organized along a continuum ranging from (1) those in which nitrification and denitrification are tightly coupled in space and time to (2) those in which nitrate production and denitrification are relatively decoupled. In aquatic ecosystems, N inputs influence denitrification rates whereas hydrology and geomorphology influence the proportion of N inputs that are denitrified. Relationships between denitrification and water residence time and N load are remarkably similar across lakes, river reaches, estuaries, and continental shelves. Spatially distributed global models of denitrification suggest that continental shelf sediments account for the largest portion (44%) of total global denitrification, followed by terrestrial soils (22%) and oceanic oxygen minimum zones (OMZs; 14%). Freshwater systems (groundwater, lakes, rivers) account for about 20% and estuaries 1% of total global denitrification. Denitrification of land-based N sources is distributed somewhat differently. Within watersheds, the amount of land-based N denitrified is generally highest in terrestrial soils, with progressively smaller amounts denitrified in groundwater, rivers, lakes and reservoirs, and estuaries. A number of regional exceptions to this general trend of decreasing denitrification in a downstream direction exist, including significant denitrification in continental shelves of N from terrestrial sources. Though terrestrial soils and groundwater are responsible for much denitrification at the watershed scale, per-area denitrification rates in soils and groundwater (kg N·km −2 ·yr −1 ) are, on average, approximately one-tenth the per-area rates of denitrification in lakes, rivers, estuaries, continental shelves, or OMZs. A number of potential approaches to increase denitrification on the landscape, and thus decrease N export to sensitive coastal systems exist. However, these have not generally been widely tested for their effectiveness at scales required to significantly reduce N export at the whole watershed scale.

Ecological Applications↗

Optimizing control of a freshwater invader in time and space

The global spread of invasive species in aquatic ecosystems has prompted population control efforts to mitigate negative impacts on native species and ecosystem functions. Removal programs that optimally allocate removal effort across space and time offer promise for improving invader suppression or eradication, especially given the limited resources available to these programs. However, science-based guidance to inform such programs remains limited. This study leverages two intensive fish removal programs for nonnative green sunfish (Lepomis cyanellus) in intermittent streams of the Bill Williams River basin in Arizona, USA, to explore alternative management strategies involving variable allocation of removal effort in time and space and compare static versus dynamic decision rules. We used Bayesian hierarchical modeling to estimate demographic parameters using existing removal data, with evidence that both removal programs led to at least a 0.39 probability of eradication. Simulated alternative management strategies revealed that population suppression, but not eradication, could be achieved with reduced effort, and that dynamic management practices that respond to species abundance in real time can improve the efficiency of removal efforts. High removal frequency and program duration, including continued monitoring after zero fish were captured, contributed to successful population control. With management efforts struggling to keep pace with the rising spread and impacts of invasive species, this research demonstrates the utility of quantitative removal models to help improve invasive removal programs and robustly evaluate the success of population suppression and eradication.

Arizona↗

Predicting the impacts of Mississippi River diversions and sea-level rise on spatial patterns of eastern oyster growth rate and production

There remains much debate regarding the perceived tradeoffs of using freshwater and sediment diversions for coastal restoration in terms of balancing the need for wetland restoration versus preserving eastern oyster ( Crassostrea virginica ) production. Further complicating the issue, climate change-induced sea-level rise (SLR) and land subsidence are also expected to affect estuarine water quality. In this study, we developed a process-based numerical modeling system that couples hydrodynamic, water quality, and oyster population dynamics. We selected Breton Sound Estuary (BSE) (∼2740 km 2 ) in the eastern Mississippi River Deltaic Plain since it is home to several of the largest public oyster seed grounds and private leases for the Gulf coast. The coupled oyster population model was calibrated and validated against field observed oyster growth data. We predicted the responses of oyster population in BSE to small- (142 m 3 s −1 ) and large-scale (7080 m 3 s −1 ) river diversions at the Caernarvon Freshwater Diversion structure planned in the 2012 Coastal Master Plan (Louisiana) under low (0.38 m) and high (1.44 m) relative sea-level rise (RSLR = eustatic SLR + subsidence) compared to a baseline condition (Year 2009). Model results showed that the large-scale diversion had a stronger negative impact on oyster population dynamics via freshening of the entire estuary, resulting in reduced oyster growth rate and production than RSLR. Under the large-scale diversion, areas with optimal oyster growth rates (>15 mg ash-free dry weight (AFDW) oyster −1 wk −1 ) and production (>500 g AFDW m −2 yr −1 ) would shift seaward to the southeastern edge of the estuary, turning the estuary into a very low oyster production system. RSLR however played a greater role than the small-scale diversion on the magnitude and spatial pattern of oyster growth rate and production. RSLR would result in an overall estuary-wide decrease in oyster growth rate and production as a consequence of decreased salinities in the middle and lower estuary because rising sea level likely causes increased stage and overbank flow downstream along the lower Mississippi River.

Louisiana↗

Carbon stock trends of baldcypress knees along climate gradients of the Mississippi River Alluvial Valley using allometric methods

Carbon stock trends of the knees of Taxodium distichum likely vary across climate gradients of the southeastern United States and contribute an unknown quantity of “teal” carbon to inland freshwater wetlands. Knee metrics (e.g., density, height, biomass) were measured in mixed T. distichum swamps across the Mississippi River Alluvial Valley (MRAV) from Illinois to Louisiana. Based on their geometric similarity to a cone, the biomasses of field knees were estimated by relating the volume of their measured field dimensions to lab-measured water displacement volume and biomass via volume/mass regressions (biomass (g) = 7.2230149 + 0.292902 × volume). Knees had greater height in flooded conditions (maximum height = 163 cm; Goose Lake, Arkansas), and also in climate normal environments of mid-range precipitation and temperature (p < 0.0001). Overall, knee biomass ha−1 was 7.5 times greater in flooded vs. not flooded conditions (34.6 ± 7.3 vs. 4.6 ± 1.0 Mg ha−1, respectively). The overall mean of knee carbon biomass stock was substantial (flooded vs. not flooded conditions: 18.1 ± 3.7 Mg C ha−1 to 2.9 ± 0.7 Mg C ha−1, respectively; knee/tree live standing biomass: 17.9–5.2%, respectively). Clearly, T. distichum knees should not be ignored in blue (teal) carbon discussions of wetlands. Because knees respond to climate normal conditions, hotter/drier environments in the MRAV could lead to a decline in the contribution of knee carbon stock in the southeastern United States.

Mississippi River Alluvial Valley↗

Ecology of selected marine communities in Glacier Bay: Zooplankton, forage fish, seabirds and marine mammals

We studied oceanography (including primary production), secondary production, small schooling fish (SSF), and marine bird and mammal predators in Glacier Bay during 1999 and 2000. Results from these field efforts were combined with a review of current literature relating to the Glacier Bay environment. Since the conceptual model developed by Hale and Wright (1979) ‘changes and cycles’ continue to be the underlying theme of the Glacier Bay ecosystem. We found marked seasonality in many of the parameters that we investigated over the two years of research, and here we provide a comprehensive description of the distribution and relative abundance of a wide array of marine biota. Glacier Bay is a tidally mixed estuary that leads into basins, which stratify in summer, with the upper arms behaving as traditional estuaries. The Bay is characterized by renewal and mixing events throughout the year, and markedly higher primary production than in many neighboring southeast Alaska fjords (Hooge and Hooge, 2002). Zooplankton diversity and abundance within the upper 50 meters of the water column in Glacier Bay is similar to communities seen throughout the Gulf of Alaska. Zooplankton in the lower regions of Glacier Bay peak in abundance in late May or early June, as observed at Auke Bay and in the Gulf of Alaska. The key distinction between the lower Bay and other estuaries in the Gulf of Alaska is that a second smaller peak in densities occurs in August. The upper Bay behaved uniformly in temporal trends, peaking in July. Densities had begun to decline in August, but were still more than twice those observed in that region in May. The highest density of zooplankton observed was 17,870 organisms/m3 in Tarr Inlet during July. Trends in zooplankton community abundance and diversity within the lower Bay were distinct from upper-Glacier Bay trends. Whereas the lower Bay is strongly influenced by Gulf of Alaska processes, local processes are the strongest influence in the upper-Bay. We identified 55 species of fish during this study (1999 and 2000) from beach seines, mid-water trawls, and rod and line catches. The diversity of physical, oceanographic, and glacial chronological conditions within Glacier Bay contribute a suite of factors that influence the distribution and abundance of fish. Accordingly, we observed significant differences in the abundance and distribution of fish within the Bay. Most significantly, abundance and diversity (primarily juvenile fish including walleye Pollock, eelblennies, and capelin) were greatest at the head of both the east and west arms where zooplankton abundance was greatest – in close proximity to tidewater glaciers and freshwater runoff. All of Glacier Bay and Icy Strait were surveyed hydroacoustically for plankton and fish during June 1999 surveys. Acoustically determined forage biomass was concentrated in relatively few important areas such as Pt. Adolphus, Berg Bay, on the Geikie-Scidmore shelf, around the Beardslee/Marble islands, and the upper arms of Glacier Bay. Forage biomass (primarily small schooling fish and euphausiids) was concentrated in shallow, nearshore waters; 50 % of acoustic biomass was found at depths < 35m, 80 % of biomass at depths < 80m. During our sampling, high density patches of prey were very rare, and less than 8 % of the area surveyed in Glacier Bay contained patch densities suitable (e.g., > 0.01 fish/m 3 ) for seabirds foraging on zooplankton and small schooling fish. Less than 1 % of the area contained patches suitable (e.g., >0.1 fish/m 3 ) for whales foraging on zooplankton and small schooling fish. High-density aggregations of 0.1-10 fish/m 3 were comprised mostly of schools containing capelin, pollock, herring or euphausiids (0.1-1 kg/m 3 ). During predator surveys (1999-2000), we observed 63 species of birds and 7 species of marine mammals. Seasonal distribution and abundance of these “apex” predators was highly variable by species. Glacier Bay supports high numbers of seabirds and marine mammals that consume zooplankton and small schooling fish. Nearshore areas had higher densities of both birds and marine mammals. Several areas, such as Pt. Adolphus, Berg Bay, on the Geikie-Scidmore shelf, the Beardslee/Marble islands, and the upper arms of Glacier Bay were focal points of small schooling fish and zooplankton consuming marine birds and mammals. Comparisons between surveys and a prior study (1991) suggested that the assemblage of birds and marine mammals in the Bay is undergoing change. Most notable was a clear decline in Brachyramphus spp. murrelets while other apex species are increasing or remaining stable. It should be noted that many of the birds and mammals observed during this project, e.g. mergansers, do not forage on zooplankton and small schooling fish; rather they forage on benthic fish and sessile invertebrates. While distribution and sampling data for these marine predator species are valid, this study did not sample benthic fish and sessile invertebrates. Thus, recommendations made by this project should be interpreted as generally specific to the zooplankton/small schooling fish marine food web components of the Glacier Bay Ecosystem.

Alaska↗

Salamander occupancy in headwater stream networks

1. Stream ecosystems exhibit a highly consistent dendritic geometry in which linear habitat units intersect to create a hierarchical network of connected branches. 2. Ecological and life history traits of species living in streams, such as the potential for overland movement, may interact with this architecture to shape patterns of occupancy and response to disturbance. Specifically, large-scale habitat alteration that fragments stream networks and reduces connectivity may reduce the probability a stream is occupied by sensitive species, such as stream salamanders. 3. We collected habitat occupancy data on four species of stream salamanders in first-order (i.e. headwater) streams in undeveloped and urbanised regions of the eastern U.S.A. We then used an information-theoretic approach to test alternative models of salamander occupancy based on a priori predictions of the effects of network configuration, region and salamander life history. 4. Across all four species, we found that streams connected to other first-order streams had higher occupancy than those flowing directly into larger streams and rivers. For three of the four species, occupancy was lower in the urbanised region than in the undeveloped region. 5. These results demonstrate that the spatial configuration of stream networks within protected areas affects the occurrences of stream salamander species. We strongly encourage preservation of network connections between first-order streams in conservation planning and management decisions that may affect stream species.

Freshwater Biology↗

Intraspecific phylogeography of Lasmigona subviridis (Bivalvia: Unionidae): Conservation implications of range discontinuity

A nucleotide sequence analysis of the first internal transcribed spacer region (ITS-1) between the 5.8S and 18S ribosomal DNA genes (640 bp) and cytochrome c oxidase subunit I (COI) of mitochondrial DNA (mtDNA) (576 bp) was conducted for the freshwater bivalve Lasmigona subviridis and three congeners to determine the utility of these regions in identifying phylogeographic and phylogenetic structure. Sequence analysis of the ITS-1 region indicated a zone of discontinuity in the genetic population structure between a group of L. subviridis populations inhabiting the Susquehanna and Potomac Rivers and more southern populations. Moreover, haplotype patterns resulting from variation in the COI region suggested an absence of gene exchange between tributaries within two different river drainages, as well as between adjacent rivers systems. The authors recommend that the northern and southern populations, which are reproductively isolated and constitute evolutionarily significant lineages, be managed as separate conservation units. Results from the COI region suggest that, in some cases, unionid relocations should be avoided between tributaries of the same drainage because these populations may have been reproductively isolated for thousands of generations. Therefore, unionid bivalves distributed among discontinuous habitats (e.g. Atlantic slope drainages) potentially should be considered evolutionarily distinct. The DNA sequence divergences observed in the nuclear and mtDNA regions among the Lasmigona species were congruent, although the level of divergence in the COI region was up to three times greater. The genus Lasmigona , as represented by the four species surveyed in this study, may not be monophyletic.

Delaware, Maryland, North Carolina, Pennsylvania, ↗

Secondary contact erodes Pleistocene diversification in a wide-ranging freshwater mussel (Quadrula)

The isolated river drainages of eastern North America serve as a natural laboratory to investigate the roles of allopatry and secondary contact in the evolutionary trajectories of recently diverged lineages. Drainage divides facilitate allopatric speciation, but due to their sensitivity to climatic and geomorphological changes, neighboring rivers frequently coalesce, creating recurrent opportunities of isolation and contact throughout the history of aquatic lineages. The freshwater mussel Quadrula quadrula is widely distributed across isolated rivers of eastern North America and possesses high phenotypic and molecular variation across its range. We integrate sequence data from three genomes, including female- and male-inherited mitochondrial markers and thousands of nuclear encoded SNPs with morphology and geography to illuminate the group's divergence history. Across contemporary isolated rivers, we found continuums of molecular and morphological variation, following a pattern of isolation by distance. In contact zones, hybridization was frequent with no apparent fitness consequences, as advanced hybrids were common. Accordingly, we recognize Q. quadrula as a single cohesive species with subspecific variation ( Q. quadrula rumphiana ). Demographic modeling and divergence dating supported a divergence history characterized by allopatric vicariance followed by secondary contact, likely driven by river rearrangements and Pleistocene glacial cycles. Despite clinal range-wide variation and hybridization in contact zones, the process-based species delimitation tool delimitR, which considers demographic scenarios like secondary contact, supported the delimitation of the maximum number of species tested. As such, when interpreting species delimitation results, we suggest careful consideration of spatial sampling and subsequent geographic patterns of biological variation, particularly for wide-ranging taxa.

eastern North America↗

Characterizing mauka-to-makai connections for aquatic ecosystem conservation on Maui, Hawaiʻi

Mauka-to-makai (mountain to sea in the Hawaiian language) hydrologic connectivity – commonly referred to as ridge-to-reef – directly affects biogeochemical processes and socioecological functions across terrestrial, freshwater, and marine systems. The supply of freshwater to estuarine and nearshore environments in a ridge-to-reef system supports the food, water, and habitats utilized by marine fauna . In addition, the ecosystem services derived from this land-to-sea connectivity support social and cultural practices (hereafter referred to as socio-cultural) including fishing, aquaculture, wetland agriculture, religious ceremonies, and recreational activities. To effectively guide island resource management, a better understanding of the linkages from ridge-to-reef across natural and social usages is critical, particularly in the context of climate change, with anticipated increasing temperature and shifting precipitation patterns. The objective of this study was to identify spatial linkages that promote multiple and diverse uses, following the ridge-to-reef concept, at an island-wide scale to identify regions of high conservation importance for aquatic resources. We selected the Island of Maui as a study representative of many Pacific islands. Diverse datasets, including agricultural lands within watersheds, wetland locations, presence of stream species, indicators of freshwater input from streams, coral cover, nearshore fish biomass, socio-cultural data such as fishpond locations, wetland taro cultivation, beach recreation use, and lastly the dynamically downscaled Coupled Model Intercomparison Project Phase (CMIP5) future climate projections scenarios (Representative Concentration Pathway (RCP) 4.5 & 8.5) were used to examine the spatial linkages through hydrological connectivity from land to the sea. Zonation spatial planning software was used to prioritize areas of high management and conservation value and to help inform aquatic resources management. The resulting prioritized areas included many minimally disturbed watersheds in east Maui and western nearshore and coastal zones that are adjacent to diverse coral reefs. These results are driven by the importance of fish biomass and coral reef distribution as well as traditional wetland taro cultivation and coastal access points for recreation. These results underline the importance of examining ridge-to-reef systems for aquatic resource management and including important social and cultural values in resource management upon planning adaptation strategies for climate change. Improving our understanding of diverse natural and socio-cultural influences on habitat conditions and their values in these areas provides an opportunity to strategically plan future management and conservation actions.

Hawaii↗

Dynamics of mangrove-marsh ecotones in subtropical coastal wetlands: fire, sea-level rise, and water levels

Ecotones are areas of sharp environmental gradients between two or more homogeneous vegetation types. They are a dynamic aspect of all landscapes and are also responsive to climate change. Shifts in the position of an ecotone across a landscape can be an indication of a changing environment. In the coastal Everglades of Florida, USA, a dominant ecotone type is that of mangrove forest and marsh. However, there is a variety of plants that can form the marsh component, including sawgrass (Cladium mariscus [L.] Pohl), needlegrass rush (Juncus roemerianus Scheele), and spikerush (Eleocharis spp.). Environmental factors including water depth, soil type, and occurrence of fires vary across these ecotones, influencing their dynamics. Altered freshwater inflows from upstream and increasing sea level over the past 100 years may have also had an impact. We analyzed a time series of historical aerial photographs for a number of sites in the coastal Everglades and measured change in position of mangrove–marsh ecotones. For three sites, detailed maps were produced and the area of marsh, mangrove, and other habitats was determined for five periods spanning the years 1928 to 2004. Contrary to our initial hypothesis on fire, we found that fire did not prevent mangrove expansion into marsh areas but may in fact assist mangroves to invade some marsh habitats, especially sawgrass. Disparate patterns in mangrove–marsh change were measured at two downstream sites, both of which had multiple fires over from 1948 to 2004. No change in mangrove or marsh area was measured at one site. Mangrove area increased and marsh area decreased at the second of these fire-impacted sites. We measured a significant increase in mangrove area and a decline in marsh area at an upstream site that had little occurrence of fire. At this site, water levels have increased significantly as sea level has risen, and this has probably been a factor in the mangrove expansion.

Florida↗

Remarkable invasion of San Francisco Bay (California, USA) by the Asian clam Potamocorbula amurensis . II. Displacement of a former community

Long-term macrobenthic sampling at a site in northern San Francisco Bay has provided an unusual opportunity for documenting the time course of an invasion by a recently introduced Asian clam Potamocorbula arnurensis . Between 1977, when sampling began, and 1986, when the new clam was first discovered, the benthic community varied predictably in response to river inflow. During years of normal or high river inflow, the community consisted of a few brackish or freshwater species. During prolonged periods of low river inflow, the number of species doubled as estuarine species (e.g. Mya arenana ) migrated up the estuary. In June 1987, at the beginning of the longest dry period in recent decades, large numbers (> 12 000 m -2 ) of juvenile P. arnurensis were discovered at the site. By midsummer 1988 the new clam predominated (> 95 %) in both total number of individuals and biomass, and the expected dry-period estuarine species did not become re-established. The rapid rise of P. arnurensis to numerical dominance throughout the region of the original introduction was probably facilitated by the fact that this region of the bay had been rendered nearly depauperate by a major flood in early 1986. Once introduced, the clam had sufficient time (> 1 yr) to become well established before the salinity regime was appropriate for the return of the estuarine species. Subsequently, the new clam was apparently able to prevent the return of the dry-period community. Its ability to live in low salinity water (< 1 %o) suggests that P. amurensis may not be displaced with the return of normal winter river flow and, therefore, may have permanently changed benthic community dynamics in this region of San Francisco Bay.

Marine Ecology Progress Series↗

Hydrologic aspects of marsh ponds during winter on the Gulf Coast Chenier Plain, USA: Effects of structural marsh management

The hydrology of marsh ponds influences aquatic invertebrate and waterbird communities. Hydrologic variables in marsh ponds of the Gulf Coast Chenier Plain are potentially affected by structural marsh management (SMM: levees, water control structures and impoundments) that has been implemented since the 1950s. Assuming that SMM restricts tidal flows and drainage of rainwater, we predicted that SMM would increase water depth, and concomitantly decrease salinity and transparency in impounded marsh ponds. We also predicted that SMM would increase seasonal variability in water depth in impounded marsh ponds because of the potential incapacity of water control structures to cope with large flooding events. In addition, we predicted that SMM would decrease spatial variability in water depth. Finally, we predicted that ponds of impounded freshwater (IF), oligohaline (IO), and mesohaline (IM) marshes would be similar in water depth, temperature, dissolved oxygen ( O 2 ), and transparency. Using a priori multivariate analysis of variance (MANOVA) contrast, we tested these predictions by comparing hydrologic variables within ponds of impounded and unimpounded marshes during winters 1997-1998 to 1999-2000 on Rockefeller State Wildlife Refuge, near Grand Chenier, Louisiana. Specifically, we compared hydrologic variables (1) between IM and unimpounded mesohaline marsh ponds (UM); and (2) among IF, IO, and IM marshes ponds. As predicted, water depth was higher and salinity and O 2 were lower in IM than in UM marsh ponds. However, temperature and transparency did not differ between IM and UM marsh ponds. Water depth varied more among months in IM marsh ponds than within those of UM marshes, and variances among and within ponds were lower in IM than UM marshes. Finally, all hydrologic variables, except salinity, were similar among IF, IO, and IM marsh ponds. Hydrologic changes within marsh ponds due to SMM should (1) promote benthic invertebrate taxa that tolerate low levels of O 2 and salinity; (2) deter waterbird species that cannot cope with increased water levels; and (3) reduce waterbird species diversity by decreasing spatial variability in water depth among and within marsh ponds.

Louisiana↗

Use of Numerical Models to Simulate Transport of Sewage-Derived Nitrate in a Coastal Aquifer, Central and Western Cape Cod, Massachusetts

The unconsolidated glacial sediments underlying Cape Cod, Massachusetts compose a regional aquifer system that is used both as a source of drinking water and as a disposal site for wastewater; in addition, the discharge of clean ground water from the aquifer system is needed for the maintenance of freshwater and marine ecosystems throughout the region. Because these uses of the aquifer conflict with one another in many areas of the Cape, local and regional planners have begun to develop sustainable wastewater plans that will facilitate the disposal of wastewater while protecting water supplies and improving the health of aquatic ecosystems. To assist local and regional planners in these efforts, the U.S. Geological Survey conducted a 2-year investigation to (1) assist local and regional planners in the evaluation of potential wastewater scenarios, (2) use results and interpretation from these analyses to develop hydrologic concepts transferable throughout the region, and (3) establish and test methods that would be of use in future evaluations. Wastewater-disposal scenarios need to be evaluated in the context of the regional ground-water-flow system. For a given rate of disposal, wastewater from sites at or near a regional ground-water divide is transported in a wider arc of flow directions, flows deeper in the system, and contaminates a larger part of the aquifer than does wastewater discharged from sites farther from the divide. Also, traveltimes of wastewater from sites near a ground-water divide to receptors are longer (as much as several hundred years) than traveltimes from sites farther from the divide. Thus, wastewater disposal at or near a divide will affect a larger part of the aquifer and likely contribute wastewater to more receptors than wastewater disposal farther from a divide; however, longer traveltimes could allow for more attenuation of wastewater-derived nitrate from those sites. Ground-water-flow models and particle tracking can be used to identify advective-transport patterns downgradient from wastewater-disposal sites and estimate traveltimes; however, these tools cannot predict the distribution of mass or concentrations of wastewater constituents, such as nitrate, in the aquifer. Flow-based particle-tracking analyses can be used to estimate mass-loading rates and time-varying concentrations at wells and ecological receptors by the accounting of mass-weighted particles discharging into the receptor of interest. This method requires no additional development beyond the flow model; however, post-modeling analyses are required. In addition, the method is based on the assumption that no mass is lost during transport, an assumption that likely is not valid in many systems. Solute-transport models simulate the subsurface transport of nitrate through the aquifer and predict the distribution of the mass of a solute in the aquifer at different transport times. This method does require additional model development beyond the flow model, but can predict timevarying concentrations at receptors. Estimates of mass-loading rates require minimal post-modeling analyses. Time-varying concentrations and mass-loading rates calculated for wells in eastern Barnstable by the two methods generally were in reasonable agreement. Inherent in the flow-based particle-tracking method is the assumption that mass is conserved along a given flow line and that there is no spreading of mass in the aquifer. Although the solute-transport models also incorporate a system-wide conservation of mass, these models allow for a spreading of mass in the aquifer, and mass is not conserved along a given flow line. As a result, estimates of concentrations and mass loading rates generally were higher in particle-tracking analyses than in solute-transport simulations. Results from the two types of simulations agreed best for wells that receive large amounts of wastewater with short traveltimes (less than 10 years) because insufficient transport

Scientific Investigations Report↗

Overwintering behavior of juvenile sea turtles at a temperate foraging ground

Most freshwater and terrestrial turtle species that inhabit temperate environments hibernate to survive extreme cold periods. However, for sea turtles, the question of whether these species use hibernation as an overwintering strategy has not been resolved (Ultsch 2006 ). Felger et al. ( 1976 ) suggested that sea turtles bury themselves in mud on the seafloor and remain dormant throughout the winter, presumably not surfacing during that time. Additional researchers have described sea turtles in temperatures <15°C as lethargic, mud‐covered and buried in bottom sediment (Carr et al. 1980 , Mendonça 1983 , Ogren and McVea 1995 ). However, more recent studies suggest that sea turtles may not be as dormant in cold temperatures as previously suggested (Hochscheid et al. 2007 ). Resolving this question is difficult due to the unpredictability of winter weather patterns and the cost of advanced tracking tools required to assess these fine‐scale behaviors. However, in January 2018, unusually calm and clear marine conditions coupled with exceptionally cold weather provided us the opportunity to observe and film turtle behavior at a foraging ground in the northern Gulf of Mexico. These images, combined with previously gathered data from vessel‐based surveys and water temperature loggers, have enabled us to piece together one of the most comprehensive views of sea turtle overwintering behavior to date.

Ecology↗

Common carp disrupt ecosystem structure and function through middle-out effects

Middle-out effects or a combination of top-down and bottom-up processes create many theoretical and empirical challenges in the realm of trophic ecology. We propose using specific autecology or species trait (i.e. behavioural) information to help explain and understand trophic dynamics that may involve complicated and non-unidirectional trophic interactions. The common carp ( Cyprinus carpio ) served as our model species for whole-lake observational and experimental studies; four trophic levels were measured to assess common carp-mediated middle-out effects across multiple lakes. We hypothesised that common carp could influence aquatic ecosystems through multiple pathways (i.e. abiotic and biotic foraging, early life feeding, nutrient). Both studies revealed most trophic levels were affected by common carp, highlighting strong middle-out effects likely caused by common carp foraging activities and abiotic influence (i.e. sediment resuspension). The loss of water transparency, submersed vegetation and a shift in zooplankton dynamics were the strongest effects. Trophic levels furthest from direct pathway effects were also affected (fish life history traits). The present study demonstrates that common carp can exert substantial effects on ecosystem structure and function. Species capable of middle-out effects can greatly modify communities through a variety of available pathways and are not confined to traditional top-down or bottom-up processes.

Marine and Freshwater Research↗

Differences in impacts of Hurricane Sandy on freshwater swamps on the Delmarva Peninsula, Mid−Atlantic Coast, USA

Hurricane wind and surge may have different influences on the subsequent composition of forests. During Hurricane Sandy, while damaging winds were highest near landfall in New Jersey, inundation occurred along the entire eastern seaboard from Georgia to Maine. In this study, a comparison of damage from salinity intrusion vs. wind/surge was recorded in swamps of the Delmarva Peninsula along the Pocomoke (MD) and Nanticoke (DE) Rivers, south of the most intense wind damage. Hickory Point Cypress Swamp (Hickory) was closest to the Chesapeake Bay and may have been subjected to a salinity surge as evidenced by elevated salinity levels at a gage upstream of this swamp (storm salinity = 13.1 ppt at Nassawango Creek, Snow Hill, Maryland). After Hurricane Sandy, 8% of the standing trees died at Hickory including Acer rubrum, Amelanchier laevis, Ilex spp., and Taxodium distichum. In Plot 2 of Hickory, 25% of the standing trees were dead, and soil salinity levels were the highest recorded in the study. The most important variables related to structural tree damage were soil salinity and proximity to the Atlantic coast as based on Stepwise Regression and NMDS procedures. Wind damage was mostly restricted to broken branches although tipped&minus;up trees were found at Hickory, Whiton and Porter (species: Liquidamabar styraciflua, Pinus taeda, Populus deltoides, Quercus pagoda and Ilex spp.). These trees fell mostly in an east or east&minus;southeast direction (88o&minus;107o) in keeping with the wind direction of Hurricane Sandy on the Delmarva Peninsula. Coastal restoration and management can be informed by the specific differences in hurricane damage to vegetation by salt versus wind.

Maryland↗

Principles for urban stormwater management to protect stream ecosystems

Urban stormwater runoff is a critical source of degradation to stream ecosystems globally. Despite broad appreciation by stream ecologists of negative effects of stormwater runoff, stormwater management objectives still typically center on flood and pollution mitigation without an explicit focus on altered hydrology. Resulting management approaches are unlikely to protect the ecological structure and function of streams adequately. We present critical elements of stormwater management necessary for protecting stream ecosystems through 5 principles intended to be broadly applicable to all urban landscapes that drain to a receiving stream: 1) the ecosystems to be protected and a target ecological state should be explicitly identified; 2) the postdevelopment balance of evapotranspiration, stream flow, and infiltration should mimic the predevelopment balance, which typically requires keeping significant runoff volume from reaching the stream; 3) stormwater control measures (SCMs) should deliver flow regimes that mimic the predevelopment regime in quality and quantity; 4) SCMs should have capacity to store rain events for all storms that would not have produced widespread surface runoff in a predevelopment state, thereby avoiding increased frequency of disturbance to biota; and 5) SCMs should be applied to all impervious surfaces in the catchment of the target stream. These principles present a range of technical and social challenges. Existing infrastructural, institutional, or governance contexts often prevent application of the principles to the degree necessary to achieve effective protection or restoration, but significant potential exists for multiple co-benefits from SCM technologies (e.g., water supply and climate-change adaptation) that may remove barriers to implementation. Our set of ideal principles for stream protection is intended as a guide for innovators who seek to develop new approaches to stormwater management rather than accept seemingly insurmountable historical constraints, which guarantee future, ongoing degradation.

Freshwater Science↗

Response curves for phosphorus plume lengths from reactive-solute-transport simulations of onland disposal of wastewater in noncarbonate sand and gravel aquifers

Surface-water resources in Massachusetts often are affected by eutrophication, excessive plant growth, which has resulted in impaired use for a majority of the freshwater ponds and lakes and a substantial number of river-miles in the State. Because supply of phosphorus usually is limiting to plant growth in freshwater systems, control of phosphorus input to surface waters is critical to solving the impairment problem. Wastewater is a substantial source of phosphorus for surface water, and removal of phosphorus before disposal may be necessary. Wastewater disposed onland by infiltration loses phosphorus from the dissolved phase during transport through the subsurface and may be an effective disposal method; quantification of the phosphorus loss can be simulated to determine disposal feasibility. In 2003, the U.S. Geological Survey, in cooperation with the Massachusetts Department of Environmental Protection, initiated a project to simulate distance of phosphorus transport in the subsurface for plausible conditions of onland wastewater disposal and subsurface properties. A coupled one-dimensional unsaturated-zone and three-dimensional saturated-zone reactive-solute-transport model (PHAST) was used to simulate lengths of phosphorus plumes. Knowledge of phosphorus plume length could facilitate estimates of setback distances for wastewater-infiltration sites from surface water that would be sufficient to protect the surface water from eutrophication caused by phosphorus transport through the subsurface and ultimate discharge to surface water. The reactive-solute-transport model PHAST was used to simulate ground-water flow, solute transport, equilibrium chemistry for dissolved and sorbed species, and kinetic regulation of organic carbon decomposition and phosphate mineral formation. The phosphorus plume length was defined for the simulations as the maximum extent of the contour for the 0.015 milligram-per-liter concentration of dissolved phosphorus downgradient from the infiltration bed after disposal cessation. Duration of disposal before cessation was assumed to be 50 years into an infiltration bed of 20,000 square feet at the rate of 3 gallons per square foot per day. Time for the maximum extent of the phosphorus plume to develop is on the order of 100 years after disposal cessation. Simulations indicated that phosphorus transport beyond the extent of the 0.015 milligram-per-liter concentration contour was never more than 0.18 kilogram per year, an amount that would likely not alter the ecology of most surface water. Simulations of phosphorus plume lengths were summarized in a series of response curves. Simulated plume lengths ranged from 200 feet for low phosphorus-concentration effluents (0.25 milligram per liter) and thick (50 feet) unsaturated zones to 3,400 feet for high phosphorus-concentration effluents (14 milligrams per liter) discharged directly into the aquifer (unsaturated-zone thickness of 0 feet). Plume length was nearly independent of unsaturated-zone thickness at phosphorus concentrations in the wastewater that were less than 2 milligrams per liter because little or no phosphorus mineral formed at low phosphorus concentrations. For effluents of high phosphorus concentration, plume length varied from 3,400 feet for unsaturated-zone thickness of 0 to 2,550 feet for unsaturated-zone thickness of 50 feet. Model treatments of flow and equilibrium-controlled chemistry likely were more accurate than rates of kinetically controlled reactions, notably precipitation of iron-phosphate minerals; the kinetics of such reactions are less well known and thus less well defined in the model. Sensitivity analysis indicated that many chemical and physical aquifer properties, such as hydraulic gradient and model width, did not affect the simulated plume length appreciably, but duration of discharge, size of infiltration bed, amount of dispersion, and number of sorption sites on the aquifer sediments did affect plume length appreciably. Because simulation of plume length in carbonate-mineral sediments indicated that the plume would be substantially longer than in noncarbonate-mineral sediments, the application of the response curves in locations with carbonate-mineral sediments would be inappropriate. The effect of carbonate minerals in sediments is to increase pH, which causes decreased sorption of phosphorus on aquifer sediments. Phosphorus removal from solution by precipitation onto aquifer sediments is more efficient at high concentrations of disposed phosphorus than at low concentrations. At very low phosphorus concentrations, the solubility product of phosphorus minerals is not exceeded and no phosphorus mineral forms. An important consequence is that removal of dissolved phosphorus from the plume by processes in the subsurface is decreased the more that removal efforts are applied in treatment before wastewater is disposed. Model simulations indicate that removal of phosphorus from wastewater disposed through septic systems would have the advantage of efficient phosphorus removal in the subsurface because phosphorus concentrations are high in septic-system effluent. Short plume lengths result from wastewater disposal through septic systems because of the efficient phosphorus removal and because of the low volume of wastewater involved. The simulation results for small-volume systems are not quantitative, however, because wastewater-infiltration rates are much lower than those of the higher-volume system that was used to calibrate the model and to create the plume-length response curves. The response curves for phosphorus plume lengths, as defined by the maximum extent of the 0.015 milligram-per-liter concentration contour, is clearly defined in the model simulations, although the relation between simulated plume length and protective setback distance is subject to interpretation. Phosphorus does move beyond the point at which the simulated 0.015 milligram-per-liter concentration contour has stopped, so that a determination of protective setback distance must include a consideration of whether that continued flux, or some other flux amount, is appropriate. Also, simulations indicate that phosphorus plumes do not reach their full extent until 50 to 200 years after disposal cessation, depending on concentration of phosphorus disposed. No phosphorus plume has been monitored for that long after cessation, so there is no way to verify the long-term simulation results.

Massachusetts↗