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Effects of food deprivation on the larvae of two flatfishes

For greatest survival, first-feeding halibut Paralichthys californicus and diamond turbot Hypsopsetta guttulata required food by the day of total yolk absorption. Some halibut larvae survived if fed 1 or 2 d after yolk depletion, but their growth rate was significantly less than larvae fed earlier. Survival of 3-wk-old larvae was greater in treatments with shorter starvation periods. A small percentage of 3-wk-old halibut larvae recovered from a maximum starvation period of 4 d while 3-wk-old diamond turbot successfully resumed feeding any time during food deprivation intervals lasting up to 9 d. Longer periods of starvation resulted in significant morphological differences – diamond turbot starved longer were not only smaller, but also less developed. In the field, larvae may experience varying periods of food deprivation due to differing spatial and temporal prey patch distributions. Our results demonstrate that differences in starvation resistance, and possibly mortality under patchy feeding conditions, are ontogenetic and species-specific.

Marine Ecology Progress Series

Time-courses in the retention of food material in the bivalves Potamocorbula amurensis and Macoma balthica significance to the absorption of carbon and chromium

Time courses for ingestion, retention and release via feces of microbial food was investigated using 2 bivalves with different feeding strategies, Potamocorbula amurensis and Macoma balthica . The results showed 2 pathways for the uptake of food material in these clams. The first is represented by an initial label pulse in the feces. The second pathway operates over longer time periods. Inert 51 Cr-labeled beads were used to determine time frames for these pathways. The first pathway, involving extracellular digestion and intestinal uptake, is relatively inefficient in the digestion of bacterial cells by P. amurensis but more efficient in M. balthica . The second pathway, involving intracellular digestion within the digestive gland of both clams, was highly efficient in absorbing bacterial carbon, and was responsible for most chromium uptake. Differences in the overall retention of microbial 51 Cr and 14 C relate not to gut-passage times but to the processing and release strategies of the food material by these 2 clams.

Marine Ecology Progress Series

Effect of adherent bacteria and bacterial extracellular polymers upon assimilation by Macoma balthica of sediment-bound Cd, Zn and Ag

Effects of adherent bacteria and bacterial extracellular polymer (exopolymer) upon uptake of particle-bound Cd, Zn and Ag by the deposit-feeding clam Macoma balthica were studied in the laboratory. Amorphous iron oxyhydroxide and unaltered and alkaline-extracted sediments were used as model particulates in separate, controlled deposit-feeding experiments. In general, amounts of metal taken up from ingested particles varied dramatically with the nature of the particle surface. Ingestion of contaminated iron oxide particles did not contribute to overall uptake of Cd and Ag in feeding clams, but accounted for 89 to 99% of total Zn uptake. Exopolymer adsorbed on iron oxide particles caused an increase in the biological availability of particle-bound metals in the order Ag>Cd>Zn, whereas adherent bacteria up to 3.2 X 10 11 g -1 had no effect upon amounts of metal taken up from ingested particulates. At the higher Cd and Ag concentrations employed (3.6 X 10 -7 M), feeding rates declined with increasing amounts of iron oxide-bound exopolymer, suggesting behavioral avoidance due to increased metal availability. Much of the Cd (57 %) taken up by clams feeding on unaltered estuarine sediments originated from particulates, even though particle/solute distribution of Cd (86%) was similar to that in experiments with iron oxide particles. Uptake of Cd from alkalineextracted sediments was insignificant, as it was from unamended iron oxide. However, addition of exopolymer (10 mgg -1 sediment) caused a restoration nn bioavailability of sediment-bound Cd.

Marine Ecology Progress Series

Persistence of an introduced mud flat community in south San Francisco Bay, California

The benthic invertebrate community inhabiting the extensive and sedimentologically homogeneous mudflats of South San Francisco Bay has demonstrated a high degree of constancy in both species composition and relative abundance among species throughout 10 yr of observation. The community, composed predominantly of introduced species with opportunistic lifestyles, is dominated numerically by Gemma gernma , Ampelisca abdita , and Streblospio benedicti . The key to the persistent co-occurrence of these species on the mudflats seems to lie in the combination of (1) the recurrence of minor disturbances of the mudflat habitat (e.g. sediment deposition/erosion, inundation by low-salinity water) on time scales comparable to that of life cycles; (2) opportunistic life history strategies (rapid maturity, brooding of young, multiple generations each year, ease of local dispersal of both juveniles and adults) that permit continued colonization of the mudflat surface or rapid recolonization after disturbances. Only 1 of the 3 numerically-dominant species. A. abdita , displays an annual periodicity in abundance. S. benedicti and G. gemma , through broadly flexible reproductive strategies permitted in the mild San Francisco climate, can exhibit strong recruitment at any time between spring and autumn. The most extreme community changes, involving temporary reduction or elimination of normally dominant populations, occurred as a result of anomalous disturbances such as unusual buildup and decay of an algal mat during 1 summer and prolonged periods of unusually high freshwater inflow during 2 successive winters. The introduced opportunists routinely co-occur at high densities. However, one of these, the tube-dwelling amphipod A. abdita , may control the abundance of the native mollusk Macoma balthica .

Marine Ecology Progress Series

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

Scale-dependent correlation of seabirds with schooling fish in a coastal ecosystem

The distribution of piscivorous seabirds relative to schooling fish was investigated by repeated censusing of 2 intersecting transects in the Avalon Channel, which carries the Labrador Current southward along the east coast of Newfoundland. Murres (primarily common murres Uria aalge ), Atlantic puffins Fratercula arctica , and schooling fish (primarily capelin Mallotus villosus ) were highly aggregated at spatial scales ranging from 0.25 to 15 km. Patchiness of murres, puffins and schooling fish was scale-dependent, as indicated by significantly higher variance-to-mean ratios at large measurement distances than at the minimum distance, 0.25 km. Patch scale of puffins ranged from 2.5 to 15 km, of murres from 3 to 8.75 km, and of schooling fish from 1.25 to 15 km. Patch scale of birds and schooling fish was similar m 6 out of 9 comparisons. Correlation between seabirds and schooling birds was significant at the minimum measurement distance in 6 out of 12 comparisons. Correlation was scale-dependent, as indicated by significantly higher coefficients at large measurement distances than at the minimum distance. Tracking scale, as indicated by the maximum significant correlation between birds and schooling fish, ranged from 2 to 6 km. Our analysis showed that extended aggregations of seabirds are associated with extended aggregations of schooling fish and that correlation of these marine carnivores with their prey is scale-dependent.

Newfoundland

An empirical model for estimating phytoplankton productivity in estuaries

e have previously shown that primary productivity in San Francisco Bay, USA, is highly correlated with phytoplankton biomass B (chlorophyll a concentration) and an index of light avallability in the photic zone, 2, I, (photic depth times surface irradiance). To test the generality of this relation, we compiled data from San Francisco Bay and 5 other USA estuarine systems (Neuse and South Rivers, Puget Sound, Delaware Bay and Hudson River Plume), and regressed daily produclvity J' P (mg C m-2 d-') against the composite parameter B Z, I,. Regressions for each estuary were significant and typically over 80 % of the varialon in P was correlated with variations in B Z,I,. Moreover, the pooled data (n = 211) from 4 estuaries where methodologies were comparable fell along one regression line (r2= 0.82), indicating that primary productivity can be estimated in a diversity of estuarine waters from simple measures of phytoplankton biomass and hght availability. This implies that physiological variabhty (e. g. responses to variations in nutrient availabhty, temperature, sahnity, photoperiod) is a secondary control on phytoplankton production in nutrient-rich estuaries, and that one empirical function can be used to estimate seasonal variations in productivity or to map productivity along estuarine gradients of phytoplankton biomass and turbidity.

Marine Ecology Progress Series

Significance of biomass and light availability to phytoplankton productivity in San Francisco Bay

Primary productivity was measured monthly at 6 sites within San Francisco Bay, USA, throughout 1980. The 6 sites were chosen to represent a range of estuarine environments with respect to salinity, phytoplankton community composition, turbidity, and water depth. Annual net production over the photic zone ranged from 95 to 150 g C m-2, and was highest in regions of lowest turbidity. Daily photic zone net productivity PN,, ranged from 0.05 to 2.2 g C m-2 d-', and was significantly correlated with the composite parameter B I,/& (where B = phytoplankton biomass; I, = daily surface insolation; E = attenuation coefficient). Lnear regression of PN,, against B Io/€ indicated that most (82 %) of the spatio-temporal variability in primary productivity within this estuary is explained by variations in light availability and phytoplankton biomass. We also calculated annual water-column net productivity PN, as a fraction of annual gross productivity PGx The ratio PN,, : PG, was inversely related to the ratio of water depth H to annual mean photic depth Z,. This linear relation indicates that the watercolumn of San Francisco Bay is a net photosynthetic source of organic carbon only when the ratio H : Z, < 6. In deep turbid habitats, where H : Tp > 6, respiratory loss exceeds productivity. Thus, 2 empirical formulations allow us to estimate productivity over the photic zone and water column from simple properties that are easily measured.

California

From understanding to action: Integrating new and old methodologies to manage marine infectious disease

Marine diseases can have far-reaching effects on population, community and ecosystem health; however, our ability to track, predict and manage these diseases has, historically, been poor. As a result, the fields of disease ecology and epidemiology have developed at a slower pace for marine than terrestrial systems [ 1 ]. New methodologies, including genomic tools for diagnostics [ 2 , 3 ], transcriptomic tools for measuring host and pathogen responses to infection (e.g. [ 4 , 5 ]), regional oceanic modelling systems that estimate environmental conditions influencing pathogen dispersal and disease progression [ 6 ], artificial intelligence methods for quantifying pathology from images (e.g. [ 7 ]) and advanced disease modelling techniques [ 8 , 9 ] are precipitating a rapid increase in our understanding of marine pathosystems. In 2016, these efforts led to the first special issue of Philosophical Transactions of the Royal Society B ( Marine diseases, volume 371, issue 1689) focused entirely on marine disease ecology and evolution, and in 2020, the first book, Marine disease ecology, was devoted to this topic [ 10 ]. This special issue, focused on marine disease management , is being published a decade after the first Philosophical Transactions special issue on marine diseases. The shift to a management focus reflects an urgent need for management strategies to address high-impact diseases and the rapid methodological advances that have resulted. The papers included in this issue demonstrate the value of combining classical approaches (e.g. routine disease surveillance, reductionistic pathogen challenge trials, rapid throughput diagnostics) with cutting-edge technologies (e.g. high-resolution oceanographic models, Bayesian models, replicated transcriptomic studies) to identify drivers of disease, quantify impacts and suggest management strategies.

Philosophical Transactions of the Royal Society, S

Using advanced population genomics to better understand the relationship between offshore and spawning habitat use for Atlantic Sturgeon

Atlantic Sturgeon ( Acipenser oxyrinchus oxyrinchus ) are a large-bodied anadromous fish that historically supported important fisheries along the east coast of the United States. Following years of overharvest and habitat degradation, populations experienced severe declines. In 2012, the National Marine Fisheries Service listed Atlantic Sturgeon under the Endangered Species Act (ESA; 61 FR 4722). Their listing named five Distinct Population Segments (DPSs), predicated on genetic groups composed of geographically proximate populations. Federal management of Atlantic Sturgeon presents challenges, as sturgeon from each of the five DPSs mix extensively in coastal and marine habitats yet take and recovery progress must be evaluated separately for each unit. Genetic assignment testing based on mitochondrial and microsatellite markers allows individuals to be assigned back to their natal river and DPS. However, this approach is not perfect and some individuals may be incorrectly assigned. Recent advances in genomics offer the potential of a higher resolution approach to genetic assignment testing, and thus may reduce uncertainty associated with assignment testing. In addition, genomics allows a greater number of markers to be examined from across a broader portion of the sturgeon genome, thus may provide an enhanced perspective of population structure for the species, and potentially allow other previously intractable questions to be addressed (Bernatchez et al. 2017, Supple and Shapiro 2018). We used next-generation sequencing to develop a draft genome for Atlantic Sturgeon and identify single nucleotide polymorphisms (SNPs) that could be used to resolve the natal river and DPS of individual Atlantic Sturgeon. We identified 1,210 candidate SNPs within the nuclear genome as well as 49 SNPs within the mitochondrial genome. After filtering and review, we selected 161 nuclear SNPs and 39 mitochondrial SNPs for further testing and evaluation. We used genotyping-in-thousands by sequencing (GT-seq) to simultaneously sequence nuclear SNP loci, mitochondrial SNP loci, and the existing panel of twelve microsatellite loci. This effort required a pilot sequencing run on a single sturgeon sample to test marker amplification and refine primer strengths, followed by a series of sequencing runs to generate baseline data for 288 individuals representing nine populations of Atlantic Sturgeon in four DPSs. Using baseline data from the nine populations, we ran a series of genomic analyses to characterize diversity within and among populations, providing a benchmark for this species using the new SNP markers. Allelic richness was similar for all populations, although there was a general trend of more northern population containing greater levels of allelic richness. Interestingly, we observed linkage disequilibrium among many pairs of loci within many populations. This might be the result of physical linkage but could also suggest these populations are recovering from genetic bottlenecks and/or are effectively small, leading to specific haplotypes to be favored by chance. Pairwise differentiation among populations varied among the populations ( F ST range: 0.010-0.098) and was significantly correlated ( r = 0.771; P < 0.001) to pairwise F ST observed using microsatellite markers). Population clustering and ordination techniques using the new genomic data both support an overall population structure that is similar to the current DPS management units (which were developed primarily based on microsatellite genetic data). Overall, this suggests that existing microsatellite markers and the panel of SNP markers developed in this study provide similar information about the populations structure and ecology of Atlantic Sturgeon. Given the observed differences in allele frequencies among populations, our genomic baseline supports previous assertations that Atlantic Sturgeon show natal homing, despite mixing extensively in marine waters during non-breeding periods. Lower levels of differentiation between populations in the South Atlantic DPS suggest that populations in this region may have greater levels of gene flow relative to their more northerly conspecifics, which has also previously been suggested based on microsatellite data. The observed differentiation among populations provides the necessary foundation for determining the natal river and DPS of Atlantic Sturgeon using assignment testing. We tested the utility of our new genomic baseline for resolving the population and DPS of Atlantic Sturgeon. Our nuclear SNP markers showed utility for identifying the origin of unknown Atlantic Sturgeon samples, as 86.5% were assigned to the correct DPS and 66.3% were assigned to the correct natal river. However, since this study was funded the Conservation Genetics and Genomics Laboratory at Leetown Science Center has made significant improvements to their microsatellite genetic baseline, which now performs more effectively than our new genomic approach (the genetic baseline includes 12 populations and 5 DPSs, and correctly assigns 95.8% of individuals to DPS and 84.9% of individuals to their natal population using 12 microsatellite loci). We conducted an ad hoc exploration of how additional microsatellite or nuclear SNP loci may further improve the accuracy of assignment testing. We found that additional microsatellite markers are likely to result in greater improvements in assignment efficiency than additional nuclear SNPs. However, a much larger number of SNP loci (which if identified could be sequenced using other methods that are now available; e.g., the RAD-capture approach published by Ali et al. 2016) could produce assignment efficiencies that are greater than what is currently feasible using microsatellites. In the absence of further research and development of additional SNP markers for Atlantic Sturgeon (possibly using an approach other than GT-seq), the existing microsatellite loci are the most effective means available to determine the natal river and DPS of Atlantic Sturgeon encountered in offshore waters. Because our new genomic markers were less effective than the existing panel of 12 microsatellite markers, we chose to use the existing microsatellite markers to assign Atlantic Sturgeon captured in another BOEM-funded study (cooperative agreement M16AC00003; Monitoring endangered Atlantic Sturgeon and commercial finfish habitat use offshore New York) following consultation with our project officer. Using this approach, we genotyped and assigned 186 Atlantic Sturgeon captured in coastal waters off the Rockaway Peninsula, New York. The vast majority of these sturgeon were assigned to the New York Bight DPS (94.62%), and most appear to belong to the Hudson River population (87.10%) with smaller contributions from the Delaware River population (7.53%). Smaller contributions (2.15%) were observed from six other populations, including those from the James, York, Kennebec, Ogeechee, and Edisto rivers. Although most of the fish we assigned were assigned to the nearest spawning rivers (Hudson and Delaware), the contributions from distant rivers is consistent with the propensity of this species to move long distances and form mixed stock aggregations along the continental shelf. This finding indicates that spawning populations (and their corresponding DPS) from distant locations may potentially be impacted by offshore activities. In fact, activities in this region of the New York Bight could negatively impact Atlantic Sturgeon population from at least four different DPSs. Genetic or genomic assignment testing remains an essential tool to characterize potential impacts to Atlantic Sturgeon populations and should be applied more broadly to better characterize potential impacts of activities in other locations.

Atlantic Coast

Proceedings of the Fourth Glacier Bay Science Symposium

Foreword Glacier Bay was established as a National Monument in 1925, in part to protect its unique character and natural beauty, but also to create a natural laboratory to examine evolution of the glacial landscape. Today, Glacier Bay National Park and Preserve is still a place of profound natural beauty and dynamic landscapes. It also remains a focal point for scientific research and includes continuing observations begun decades ago of glacial processes and terrestrial ecosystems. In recent years, research has focused on glacial-marine interactions and ecosystem processes that occur below the surface of the bay. In October 2004, Glacier Bay National Park convened the fourth in a series of science symposiums to provide an opportunity for researchers, managers, interpreters, educators, students and the general public to share knowledge about Glacier Bay. The Fourth Glacier Bay Science Symposium was held in Juneau, Alaska, rather than at the Park, reflecting a desire to maximize attendance and communication among a growing and diverse number of stakeholders interested in science in the park. More than 400 people attended the symposium. Participants provided 46 oral presentations and 41 posters covering a wide array of disciplines including geology, glaciology, oceanography, wildlife and fisheries biology, terrestrial and marine ecology, socio-cultural research and management issues. A panel discussion focused on the importance of connectivity in Glacier Bay research, and keynote speakers (Gary Davis and Terry Chapin) spoke of long-term monitoring and ecological processes. These proceedings include 56 papers from the symposium. A summary of the Glacier Bay Science Plan-itself a subject of a meeting during the symposium and the result of ongoing discussions between scientists and resource managers-also is provided. We hope these proceedings illustrate the diversity of completed and ongoing scientific studies, conducted within the Park. To this end, we invited all presenters to submit brief technical summaries of their work, to capture the gist of their study and its main findings without an overload of details and methodology. We also asked authors to include a few words on the management implications of their work to help bridge the gap between scientists and managers in understanding how specific research questions may translate to management practice. Papers in this volume are laid out by subject matter, from terrestrial and freshwater subjects to glacial-marine geology, to the ecology of marine animals and ending with risk assessment, human impacts and science-management considerations. In summary, we hope the proceedings will serve as a useful reference to completed and ongoing studies in Glacier Bay National Park, and thereby provide park enthusiasts, scientists, and managers with a road map of scientific progress.

Scientific Investigations Report

Land-margin ecosystem hydrologic data for the coastal Everglades, Florida, water years 1996-2012

Mangrove forests and salt marshes dominate the landscape of the coastal Everglades (Odum and McIvor, 1990). However, the ecological effects from potential sea-level rise and increased water flows from planned freshwater Everglades restoration on these coastal systems are poorly understood. The National Park Service (NPS) proposed the South Florida Global Climate Change Project (SOFL-GCC) in 1990 to evaluate climate change and the effect from rising sea levels on the coastal Everglades, particularly at the marsh/mangrove interface or ecotone (Soukup and others, 1990). A primary objective of SOFL-GCC project was to monitor and synthesize the hydrodynamics of the coastal Everglades from the upstream freshwater marsh to the downstream estuary mangrove. Two related hypotheses were set forward (Nuttle and Cosby, 1993): 1. There exists hydrologic conditions (tide, local rainfall, and upstream water deliveries), which characterize the location of the marsh/mangrove ecotone along the marine and terrestrial hydrologic gradient; and 2. The marsh/mangrove ecotone is sensitive to fluctuations in sea level and freshwater inflow from inland areas. Hydrologic monitoring of the SOFL-GCC network began in 1995 after startup delays from Hurricane Andrew (August 1992) and organizational transfers from the NPS to the National Biological Survey (October 1993) and the merger with the U.S. Geological Survey (USGS) Biological Research Division in 1996 (Smith, 2004). As the SOFL-GCC project progressed, concern by environmental scientists and land managers over how the diversion of water from Everglades National Park would affect the restoration of the greater Everglades ecosystem. Everglades restoration scenarios were based on hydrodynamic models, none of which included the coastal zone (Fennema and others, 1994). Modeling efforts were expanded to include the Everglades coastal zone (Schaffranek and others, 2001) with SOFL-GCC hydrologic data assisting the ecological modeling needs. In 2002, as a response for a more interdisciplinary science approach to understanding the coastal Everglades ecological system, the SOFL-GCC hydrology project was integrated into the “Dynamics of Land-Margin Ecosystems: Historical Change, Hydrology, Vegetation, Sediment, and Climate” study (Smith and others, 2002). Data from the ongoing study has been useful in providing an empirical hydrologic baseline for the greater Everglades ecosystem restoration science and management needs. The hydrology network consisted of 13 hydrologic gaging stations installed in the southwestern coastal region of Everglades National Park along three transects: Shark River (Shark or SH) transect, Lostmans River (Lostmans or LO) transect, and Chatham River (Chatham or CH) transect (fig. 1). There were five paired surface-water/groundwater gaging stations on the Shark transect (SH1, SH2, SH3, SH4, and SH5) and one stage gaging station (BSC) in the Big Sable Creek; four paired surface-water/groundwater gaging stations on the Lostmans transect (LO1, LO2, LO3, and LO4); and three paired surface-water/groundwater gaging stations on the Chatham transect (CH1, CH2, and CH3). Both surface-water and groundwater levels, salinities, and temperatures were monitored at the paired gaging stations. Rainfall was recorded at marsh and open canopy gaging stations. This report details the study introduction, method, and description of data collected, which are accessible through the final instantaneous hydrologic dataset stored in the USGS South Florida Information Access (SOFIA) South Florida Hydrology Database website, http://sofia.usgs.gov/exchange/sfl_hydro_data/location.html#brdlandmargin .

Florida

Scalability and performance tradeoffs in quantifying relationships between elevation and tidal wetland plant communities

Elevation is a major driver of plant ecology and sediment dynamics in tidal wetlands, so accurate and precise spatial data are essential for assessing wetland vulnerability to sea-level rise and making forecasts. We performed survey-grade elevation and vegetation surveys of the Global Change Research Wetland, a brackish microtidal wetland in the Chesapeake Bay estuary, Maryland (USA), to both intercompare unbiased digital elevation model (DEM) creation techniques and to describe niche partitioning of several common tidal wetland plant species. We identified a tradeoff between scalability and performance in creating unbiased DEMs, with more data-intensive methods such as kriging performing better than 3 more scalable methods involving post-processing of light detection and ranging (LiDAR)-based DEMs. The LiDAR Elevation Correction with Normalized Difference Vegetation Index (LEAN) method provided a compromise between scalability and performance, although it underpredicted variability in elevation. In areas where native plants dominated, the sedge Schoenoplectus americanus occupied more frequently flooded areas (median: 0.22, 95% range: 0.09 to 0.31 m relative to North America Vertical Datum of 1988 [NAVD88]) and the grass Spartina patens , less frequently flooded (0.27, 0.1 to 0.35 m NAVD88). Non-native Phragmites australis dominated at lower elevations more than the native graminoids, but had a wide flooding tolerance, encompassing both their ranges (0.19, -0.05 to 0.36 m NAVD88). The native shrub Iva frutescens also dominated at lower elevations (0.20, 0.04 to 0.30 m NAVD88), despite being previously described as a high marsh species. These analyses not only provide valuable context for the temporally rich but spatially restricted data collected at a single well-studied site, but also provide broad insight into mapping techniques and species zonation.

Maryland