Uptake and release of phosphorus by phytoplankton in the Chesapeake Bay estuary,USA

The phytoplankton uptake and release rates for inorganic phosphate, dissolved organic phosphate and polyphosphate were estimated during 5 cruises on the Chesapeake Bay over a 9-month period. Phosphorus in all pools turned over in several minutes to 100 h, and each soluble pool appeared to contain fractions which were metabolically useful to the phytoplankton. Maximal uptake rates (V _m ) for orthophosphate ranged from 0.02 to 2.95 μg-at P (1.h)^-1 with half saturation constants (K _s ) between 0.09 and 1.72 μg-at P l^-1. At low soluble reactive phosphorus concentrations, the uptake rate of trace ³²P orthophosphate was initially rapid, but declined after 15 to 60 min incubation. The data suggest that the initial uptake phase was dominated by exchange of ³²PO₄ ^≡ for ³¹PO₄ ^≡ in the membrane transport systems whereas the subsequent phase represented the net incorporation of orthophosphate into phytoplankton cells.     

Economic losses associated with the degradation of an ecosystem: The case of submerged aquatic vegetation in Chesapeake Bay

This study employs theoretical and empirical concepts from ecology and economics to derive a lower bound of the marginal damage function for reductions in the level of submerged aquatic vegetation (SAV) in Chesapeake Bay. These reductions in SAV are believed to be a consequence of the runoff of agricultural chemicals, discharges from waste treatment plants, and soil erosion. The study examines the indirect ecological consequences of pollution in Chesapeak Bay fisheries, in a fashion which is consistent with the economic theory of benefit measurement.     

Distribution of the cladoceran Podon polyphemoides in the Chesapeake Bay

The distribution of the cladoceran Podon polyphemoides (Leuckart) in the Chesapeake Bay (USA) estuarine system was determined by a quantitative pump sampling method, and the patterns of abundance were correlated with temperature and salinity distributions. The species was seasonally recurrent, with distinct population maxima in the central portion of the bay. Population densities in excess of 60,000 podonids/m³ have been recorded. The podonids first appeared in the spring in the shallow tributaries, when water temperatures near the bottom reached 6°C. The vernal populations disappeared when summer temperatures exceeded 27°C, but reappeared in the fall as the water cooled. The species was euryhaline and eurythermal in its distribution, but the greatest concentrations were attained within relatively narrow zones of temperatures between 11^o and 26°C, and salinities between 8 and 18. The production of males, sexual females and sexual eggs occurred both in the spring and the fall between the thermal limits of 11^o and 17°C.     

Spatial and temporal variation of resource limitation in Chesapeake Bay

We report nutrient addition bioassays at 18 stations in Chesapeake Bay (USA) to assess resources limiting phytoplankton growth. Data were pooled from several sampling programs conducted from 1989 to 1994. Spatially, light and P limitation declined from low salinity regions to high salinity regions, as N limitation increased. This spatial pattern was driven primarily by freshwater inflows with high N/P and seawater inflows with low N/P. Seasonally, there was a marked progression of winter light limitation, spring P limitation, and summer N limitation at mesohaline and polyhaline stations. The seasonal pattern appeared to be caused by temperature, mixing, river discharge, and sediment P fluxes. At high salinity stations, we also observed winter N limitation (caused by DIN depletion prior to spring nitrate delivery), and at lower salinity stations there was fall P limitation (caused by reaeration of bottom sediments). At tidal fresh stations, turbidity and nutrient concentrations resulted in continuous light limitation, except at some stations in summer. Interannual decreases in light limitation and increases in N and P limitation appear to represent improvements in water quality. Received: 31 October 1997 / Accepted: 18 December 1998     

Quantitative feeding ecology of the hydromedusan Nemopsis bachei in Chesapeake Bay

We determined feeding rates of the hydromedusan Nemopsis bachei L. Agassiz in the mesohaline region of Chesapeake Bay, USA during the spring of 1989 and 1990 from gut contents, digestion rates and abundances of medusae and zooplankton. The medusae consumed primarily copepodites of Acartia tonsa, selecting against naupliar stages. The peak abundance of N. bachei medusae was in April to May, when densities averaged more than 10 m^-3. Medusa densities were similar in both years, but were greatest (maximum of 132 medusae m^-3) along a southern transect sampled only in 1990. At peak densities, N. bachei medusae consumed 30% d^-1 of the copepodite standing stocks, but they consumed <1% d^-1 at the lower densities typical of late May or early June. The predation effects were generally greater than those reported for other hydromedusan species. But even at peak predation, N. bachei medusae could not have controlled or reduced A. tonsa copepod populations, which had a production rate of 85% d^-1 at that time. Medusa feeding rates were highest at nighttime, and were correlated with prey density in the field, but not in the laboratory.Communicated by J. Grassle, New Brunswick     

Grazing by adult estuarine calanoid copepods of the Chesapeake Bay

Grazing by adult female Eurytemora affinis, Acartia tonsa and A. clausi on natural distributions of particles from the Chesapeake Bay has been investigated. During the course of a year's sampling, a wide variety of particle size-biomass distributions were observed as seasonal shifts in detritus, and over 150 algal species occurred. These distributions were grouped into 5 basic types in the analyses of feeding. All three species demonstrated similar capabilities for feeding over a broad range of particle size with selection (higher filtering rates) on larger particles and on biomass peaks. Feeding on multiple-peak distributions resulted in strong selection or tracking of each biomass peak with reduced filtering rates between peaks. Evidence is presented which suggests that the copepods first feed on large particles and then successively switch to biomass peaks of the smaller size categories. Comparisons of the feeding behavior of Eurytemora affinis and the Acartia species showing that the Acartia species have greater capabilities for taking large particles may be associated with modifications of their mouth parts for raptorial feeding. The results suggest considerable flexibility in copepod feeding behavior which cannot be explained solely by the mechanism of a fixed sieve.     

Colonization of meiobenthos in oil-contaminated subtidal sands in the lower Chesapeake Bay

In-situ manipulative experiments were conducted over a 3-month period (May–August 1980) to examine the rate at which meiobenthos colonizes oiled and untreated azoic fine sands at a shallow subtidal site in the lower York River, Virginia. Three concentrations of fresh Prudhoe Bay crude oil were added to sediments: 100, 2 500 and 10000 mg oil kg^-1 dry wt sediment. Untreated azoic and natural sediments served as controls. Within 16 d, meiofauna densities in all treatments were comparable to natural populations in surface oxidized sediments, but densities fluctuated greatly during the remainder of the sampling period. Nematodes slowly colonized the subsurface anoxic sediments below the redox potential discontinuity (RPD); some less common species did not significantly recover below the RPD in the two more heavily oiled treatments. Analysis of nematode community composition by reciprocal averaging ordination and numerical classification revealed generally lower abundances, but no distinct differences, in species composition in the oiled substrates as compared to untreated and natural community controls. Ordination of sequential samples suggested that the nematode species assemblages in the untreated controls fully recovered from these small-scale disturbances by 90 d. Life history characteritics and frequent tidal transport combine to make estuarine meiobenthos highly resilient following disturbance. Contrary to prior recolonization studies, a successional sequence was found for the colonizing nematodes which may be analogous to models of macrobenthic colonization (e.g. McCall, 1975). The comesomatid nematode Sabatieria pulchra, which is frequently dominant in polluted sediments, colonized relatively late in the experiment. Consequently, stress resistance and resilience may not be as coincident in meiofauna as in macrofauna because of differences in factors affecting their dispersal.     

Mitochondrial DNA markers allow monitoring of oyster stock enhancement in the Chesapeake Bay

Overharvesting, habitat degradation, and disease have resulted in a century of decline for Atlantic Coast populations of the eastern oyster Crassostrea virginica (Gmelin). The introduction of oysters with superior disease resistance (e.g. oysters from different geographical areas, or genetically improved strains) may be useful in restoration efforts. In 1997 the Oyster Recovery Partnership and the University of Maryland Center for Environmental Science planted more than four million Louisiana oysters in the Choptank River, which flows into the Chesapeake Bay, Maryland, USA. These oysters, which may be distinguished from Atlantic oysters by diagnostic single-nucleotide polymorphisms (SNPs) in their mitochondrial DNA, were expected to display enhanced survival and reproduction as a result of their superior resistance to Dermo disease. A high-throughput, synthesis-by-sequencing technique (Pyrosequencing) was used to determine the mitochondrial haplotypes of spat collected in the Choptank River and nearby regions of the bay. Of 3,545 spat collected in 1999, 2000, and 2001, 3,349 (94.47%) possessed the North Atlantic haplotype, 176 (4.68%) had the South Atlantic haplotype, and 3 individuals (0.08%) had the Gulf Coast haplotype. Detection of newly recruited oysters possessing the Gulf Coast haplotype in the Choptank River confirmed the survival and reproduction of the outplanted Louisiana oysters. If appropriate genetic tags are available, effective monitoring of stock-enhancement projects can be achieved with high-throughput molecular genotyping techniques.Electronic Supplementary Material Supplementary material is available in the online version of this article at Communicated by J.P. Grassle, New Brunswick     

Modeling the seasonal autochthonous sources of dissolved organic carbon and nitrogen in the upper Chesapeake Bay

In this paper we investigate the seasonal autochthonous sources of dissolved organic carbon (DOC) and nitrogen (DON) in the euphotic zone at a station in the upper Chesapeake Bay using a new mass-based ecosystem model. Important features of the model are: (1) carbon and nitrogen are incorporated by means of a set of fixed and varying C:N ratios; (2) dissolved organic matter (DOM) is separated into labile, semi-labile, and refractory pools for both C and N; (3) the production and consumption of DOM is treated in detail; and (4) seasonal observations of light, temperature, nutrients, and surface layer circulation are used to physically force the model. The model reasonably reproduces the mean observed seasonal concentrations of nutrients, DOM, plankton biomass, and chlorophyll a. The results suggest that estuarine DOM production is intricately tied to the biomass concentration, ratio, and productivity of phytoplankton, zooplankton, viruses, and bacteria. During peak spring productivity phytoplankton exudation and zooplankton sloppy feeding are the most important autochthonous sources of DOM. In the summer when productivity peaks again, autochthonous sources of DOM are more diverse and, in addition to phytoplankton exudation, important ones include viral lysis and the decay of detritus. The potential importance of viral decay as a source of bioavailable DOM from within the bulk DOM pool is also discussed. The results also highlight the importance of some poorly constrained processes and parameters. Some potential improvements and remedies are suggested. Sensitivity studies on selected parameters are also reported and discussed.     

The effect of light on strobilation in the Chesapeake Bay Sea Nettle Chrysaora quinquecirrha

Previous work (Loeb, 1972) showed that strobílation could be induced in Chrysaora quinquecirrha (Desor) by subjecting polyps to a period of chilling followed by warming. It is shown here, that the onset of strobilation and the rate at which this process occurs in a population of C. quinquecirrha polyps is positively related to the amount of light received during chilling and in the period following warming. The absence of light during either period delays the onset of strobilation. The intensity of coloration of C. quinquecirrha polyps before strobilation appears to be correlated with the amount of light to which they have been exposed during chilling.     

Grazing of the mixotrophic dinoflagellate Gymnodinium sanguineum on ciliate populations of Chesapeake Bay

In situ grazing rates for the mixotrophic dinoflagellate Gymnodinium sanguineum Hirasaka feeding on nanociliate populations of Chesapeake Bay were determined in June and October of 1990 using a gut clearance/gut fullness approach. Recently ingested prey were digested beyond the point of recognition at a rate of 23% h^-1. Estimates of in situ ingestion and clearance ranged from 0 to 0.06 prey dinoflagellate^-1 h^-1 and 0 to 5.8 l dinoflagellate^-1 h^-1, respectively, with daily removal of ciliate biomass representing 6 to 67% of the 20-m oligotrich standing stock. Daily consumption of ciliate biomass by G. sanguineum averaged 2.5% of body carbon and 4.0% of body nitrogen with maximal values of 11.6 and 18.5%, respectively. Ingestion of ciliates may help balance nitrogen requirements for G. sanguineum and give this species an advantage over purely photosynthetic dinoflagellates in nitrogen limited environments. By preying on ciliates, these dinoflagellates reverse the normal flow of material from primary producer to consumer and thereby influence trophodynamics of the microbial food web in Chesapeake Bay.     

Influence of predation on infaunal abundance in Upper Chesapeake Bay,USA

The importance of predators in controlling the densities of infaunal (>0.5 mm) organisms was investigated in the mesohaline region of the Upper Chesapeake Bay (USA) using field experiments. The role of predators in controlling infaunal density and community characteristics varied with habitat type, season (i.e., predator abundance) and developmental or successional stage of the community. Few infaunal species were adversely affected by predator exclusion. Species that increased greatly in abundance in the absence of predators (e.g. Eteone heteropoda, Streblospio benedicti, Nereis succinea, and juvenile Macoma balthica and Mya arenaria) lived near the sediment-water interface and had major population pulses from fall through spring. Species whose abundances increased moderately or were not affected by predator exclusion were deeper burrowing organisms (e.g. Heteromastus filiformis and adult Mya arenaria), or were relatively small organisms (e.g. Paraprionospio pinnata, Scolecolepides viridis and Peloscolex gabriellae) whose principal predators could be other members of the infauna. Competition did not appear to be an important factor controlling infaunal density in these experiments.This work is Contribution No. 973 of the Center for Environmental and Estuarine Studies of the University of Maryland     

Production of picoplankton and small nanoplankton in the Celtic Sea

A significant proportion of the total primary production in the Celtic Sea (50°30′N; 07°00′W) has been found to be due to picoplankton and small nanoplankton. In July, August and October, 1982, 20 to 25% of the ¹⁴C fixed in primary production was in organisms >5 μm, 35 to 40% was in organisms <5–1 μm and 20 to 30% was in organisms<1 μm. Bacterial production was estimated by the incorporation of ³H and would account for less than 10% of the production in the <1–>0.2 μm fraction; therefore, production in the <1–>0.2 μm fraction was the result of photosynthesis per se by picoplankton and could not have been due to heterotrophic bacteria utilizing exudates from larger phytoplankton. Time-course experiments demonstrated some transfer of label from the <1–>0.2μm fraction to the >5 μm fraction, presumably by grazing, but again most of the production in this fraction was the result of photosynthesis by organisms larger than 5 μm and was not due to grazing by heterotrophic microflagellates on smaller phytoplankton.     

Diurnal vertical migration of an estuarine cladoceran,Podon polyphemoides,in the Chesapeake Bay

Populations of the marine cladoceran Podon polyphemoides (Leuckart) in the Chesapeake Bay (USA) zooplankton typically occur in large patches, extending the length of the estuary. These patches are characterized by well defined maxima, which do not move seaward during a given season. During the day, the bulk of the population is situated in the upper part of the two-layered estuarine circulation, where it is subjected to transport by net non-tidal seaward current. One mechanism for maintaining the population within the estuary is provided by the diurnal behavior of the podonids, which migrate to deeper water during the night. The landward advection of the deeper currents, thus, helps to counterbalance seaward population loss during the day. The extent of the vertical movement of the population can be modified by various environmental factors, but a suggestive correlation with light was found. It is thought that the downward component of the migration is due to passive sinking.     

Mangrove forest structure and productivity in the Fly River estuary,Papua New Guinea

In April, July and August 1989 and February 1990, the delta region of the Fly River was surveyed to establish the aerial extent of mangrove forests, their species composition, tree densities and basal areas, and potential net primary production. Mangrove forests cover 87 400 ha, mainly on islands within the delta. Twentynine mangrove plant species were recorded, but there were only three major forest types in the delta.Rhizophora apiculata-Bruguiera parviflora forests (hereafterRhizophora-Bruguiera forests) predominated in regions where river water salinities were >10. These forests covered 31 500 ha and had mean total tree densities and basal areas of 2027 stems ha^–1 and 21 m² ha^–1, respectively. Forests of the palmNypa fruticans (hereafterNypa forests) covered 38 400 ha, mainly in regions where river salinities were ~1 to 10, and had mean total densities and basal areas of 4431 stems ha^–1 and 38 m² ha^–1, respectively. Forests dominated byAvicennia marina and/orSonneratia lanceolata (hereafterAvicennia-Sonneratia forests) predominated on accreting banks of sediment and covered 17 500 ha. In very low-salinity (< 1) regions there are large monospecific stands ofS. lanceolata. Mean total densities and basal areas forAvicennia-Sonneratia forests were 7036 stems ha^–1 and 22 m² ha^–1, respectively. Mean net primary productivity (kg C ha^–1 d^–1) was estimated to be 26.7, 27.1 and 19.0 forRhizophora-Bruguiera, Nypa andAvicennia-Sonneratia forests, respectively. Total daily net primary production by all mangrove forests was estimated at 2214 t carbon. Using assumptions based on work in tropical Australia, it was estimated that ~678 t carbon (or 31% of primary production) were exported daily from mangrove forests to the waters of the delta.Contribution No. 550 from the Australian Institute of Marine Science     

Ammonium excretion by gelationous zooplankton and their contribution to the ammonium requirements of microplankton in Chesapeake Bay

Ammonium excretion rates of recently collected specimens of gelatinous zooplankton, the scyphomedusan Chrysaora quinquecirrha DeSor and the etenophore Mnemiopsis leidyi A. Agassiz, were correlated with body mass and water temperature in measurements made from April to October 1989 and 1990. Rates ranged between 3.5 and 5.0 g atoms NH ₄ ⁺ -N (g dry wt)^-1h^-1 for C. quinquecirrha and 3.0 to 4.9 g atoms NH ₄ ⁺ -N (g dry wt)^-1h^-1 for M. leidyi. Excretion rate equations and in situ data on the size distributions and biomasses of gelatinous zooplankters and water temperature were used to estimate the contribution of ammonium by medusae and ctenophores to mesohaline Chesapeake Bay waters on several dates during April to October 1989 and 1990. We then compared the estimated contributions to direct measurements of ¹⁵NH ₄ ⁺ uptake by microplankton. The maximum estimated regeneration by gelatinous zooplankton was 5.8 g atoms NH ₄ ⁺ -N m^-3h^-1 at night in August 1990, when medusae biomass was greatest. This represents about 4% of the ammonium required by the microplankton. During the daytime on all dates, less than 1% of the ammonium required by microplanktion was supplied by gelatinous zooplankton. Therefore, gelatinous zooplankton appear to play a minor role in the ammonium cycle of Chesapeake Bay.     

Mortality of newly metamorphosed eastern oysters (Crassostrea virginica) in mesohaline Chesapeake Bay

 Stocks of eastern oysters, Crassostrea virginica (Gmelin), in mesohaline Chesapeake Bay, USA, exhibit a high degree of inter-annual and spatial variability in recruitment. We found that cumulative oyster spatfall on off-bottom collector plates, measured throughout the summer in 14 years over a span of three decades, was highly positively correlated (r ² = 0.8) with juvenile oyster recruitment on adjacent oyster bars. Total abundances of juvenile oysters on these bars were, however, generally 99.7% lower than predicted from cumulative seasonal larval settlement on collector plates. We propose that although the number of larvae metamorphosing was the key factor in determining the gross annual pattern of recruitment to these mesohaline oyster bars, the actual magnitude of recruitment was governed by post-settlement processes, such as competition for limited resources and predation. We tested the hypothesis that predation may be partly responsible for high post-settlement juvenile oyster mortality. We performed a series of 3-d field investigations over two summers (1989, 1990) at a mesohaline site, employing cages of various mesh sizes (400, 800, 1500 μm) to protect hatchery-reared spat of 0.5 to 4.0 mm shell height. Mortality rates for spat held for 3 d in the estuary (17.8%) were significantly higher (P = 0.0001) for the smallest spat (0.5 to 2.0 mm) compared with those of 2.01 to 4.0 mm (4.2%). In 1990, but not in 1989, enclosure within 400 and 800 μm mesh cages significantly (P = 0.004) increased survival during 3-d deployments (9.4 and 10.1%, respectively) compared with spat unprotected by mesh cages (21.9%). In a series of laboratory predation studies that used the entire community of invertebrates that could penetrate the cages, microscopic juvenile polyclad flatworms, Stylochus ellipticus, were the only organisms that we observed crawling into living oysters and feeding on oyster tissue. Large flatworms (50 to 200 mm²) are known to be important predators on oysters, but this ability of flatworms that were so small (<ca. 5 mm²) and translucent as to be almost invisible without magnification to feed on immediate post-metamorphic oysters has not been documented previously. Our results suggest that the rate of mortality due to predation in mesohaline Chesapeake Bay is much reduced once spat survive for 2 to 3 weeks post-metamorphosis. Thus, it is likely that predation in the 1 to 2 week period immediately after settlement may be a crucial factor in the structuring of eastern oyster populations. Received: 21 December 1998 / Accepted: 2 December 1999     

Mitochondrial DNA analysis of native and selectively inbred Chesapeake Bay oysters,Crassostrea virginica

Mitochondrial DNA (mtDNA) genotypes were examined in 1989 in threeCrassostrea virginica (Gmelin) populations native to Chesapeake Bay, USA, and in one population selectively inbred for rapid growth for ten generations. We wished to determine whether this character would be useful as a genetic marker for distinguishing between the inbred line and the native oysters and to determine whether detectable genetic differences exist among present-day native populations ofC. virginica. Thirty mtDNA haplotypes were identified. The average percentage nucleotide difference between native haplotypes was 1.8%. Inbred oysters were characterized by mtDNA haplotypes distinctly different from ancestral native oysters, indicating a high degree of genetic differentiation between the two groups (average percentage mtDNA nucleotide sequence divergence,=21.8%). The common native haplotype was not present in the selectively inbred sample, and six of the seven haplotypes detected in the inbred oysters were not found in the survey of native oysters. Chi-square tests on haplotype frequencies indicated that the native populations were not significantly different from one another. However, the distribution and relatedness of haplotypes suggest that significant change in the oyster gene pool may have occurred over the past few decades.     

Microphagous ciliates in mesohaline Chesapeake Bay waters: Estimates of growth rates and consumption by copepods

Growth rates of microphagous ciliates (forms which feed primarily on picoplankton-sized prey) were estimated, along with rates of their consumption by copepods, in shipboard experiments conducted in the mesohaline portion of Chesapeake Bay, USA, under contrasting water column conditions in April, June, and August 1987. Estimates were based on temporal changes in cell densities in size-fractionated water samples incubated under in situ conditions. In April, at low temperatures (7 to 10°C) and with oxygen present throughout the water column, similar generation times of ca. 1 to 1.5 d were estimated for surface and deep water (24 m) ciliate populations. In June, water was anoxic below 12 m and a distinct anoxic microphage community grew at about twice the rate of the surface community with generation times of ca. 7 and 14 h, respectively. In August, bottom water was again anoxic, but the sameStrobilidium sp. dominated both surface and deep waters with low or no growth apparent in anoxic waters and a generation time of ca. 8 h in surface waters. Copepod (primarilyAcartia tonsa Dana nauplii) clearance rates for microphagous ciliates in surface waters were 0.11, 0.56, and 0.53 ml h^–1 copepod^–1 for April, June and August, respectively. Calculation of removal rates, based on average densities, indicated that from 34 to 200% of surface waters were cleared d^–1 of microphagous ciliates by copepods.