Root respiration and oxygen flux in salt marsh grasses from different elevational zones

Plants growing in waterlogged environments are subjected to low oxygen levels around submerged tissues. While internal oxygen transport has been postulated as an important factor governing flooding tolerance, respiration rates and abilities to take up oxygen under hypoxic conditions have been largely ignored in plant studies. In this study, physiological characteristics related to internal oxygen transport, respiration, and oxygen affinity were studied in low intertidal marsh species (Spartina alterniflora and S. anglica) and middle to high intertidal species (S. densiflora, S. patens, S. foliosa, a S. alterniflora × S. foliosa hybrid, S. spartinae, and Distichlis spicata). These marsh plants were compared to the inland species S. pectinata and the crop species rice (Oryza sativa), corn (Zea mays), and oat (Avena sativa). Plants were grown in a greenhouse under simulated estuarine conditions. The low marsh species S. anglica was found to transport oxygen internally at rates up to 2.2 μmol O₂ g fresh root weight⁻¹ h⁻¹. In contrast, marsh species from higher zones and crop species were found to transport significantly less oxygen internally, although rice plants were able to transport 1.4 μmol g⁻¹ h⁻¹. Under hypoxic conditions, low marsh species were better able to remove dissolved oxygen from the medium compared to higher marsh species and crops. The oxygen concentration at which respiration rates declined due to limited oxygen (P _crit) was significantly lower in low marsh species compared to inland and crop species; P _crit ranged from <4 μM O₂ in the low marsh species S. anglica up to 20 μM in the inland species corn. Flooding-sensitive crop species had significantly higher aerobic respiration rates compared to flooding-tolerant species in this study. Crop species took up 3.6–6.7 μmol O₂ g⁻¹ h⁻¹ while all but one marsh species took up <3.5 μmol O₂ g⁻¹ h⁻¹. We conclude that oxygen transport, aerobic demand, and oxygen affinity all play important and interrelated roles in flood tolerance and salt marsh zonation.     

The ecology of temperate salt-marsh fucoids. I. Occurrence and distribution of Ascophyllum nodosum ecads

The distribution of several free-living Ascophyllum nodosum ecads, including scorpioides and mackaii, in a temperate salt marsh is described. Morphological characterization of these ecads by the presence or absence of air bladders and reproductive receptacles, and size and shape of fronds, indicated that several free-living forms occur throughout the marsh. Plants resembling the ecad mackaii were more closely associated with exposed areas along the low-tide regions, while scorpioides-type ecads prevailed on the Spartina alterniflora-dominated marsh banks and flats in the middle and upper intertidal regions of the marsh. Maximum biomass of ecads occurred during the spring months in the absence of S. alterniflora, whereas minimum ecad biomass was associated with maximum S. alterniflora densities in the late summer and fall months. Morphological differentiation of dwarf-type ecads was related to environmental components other than exposure to low and/or fluctuating salinities.This research was suported by research grant AG-375 from the National Science Foundation and, in part, by the State University of New York Research Foundation and the Energy Research and Development Administration (ERDA).     

The ecology of temperate salt-marsh fucoids. II. In situ growth of transplanted Ascophyllum nodosum ecads

Growth, in terms of length, weight, and number of branches and/or dichotomies, in transplanted specimens of Ascophyllum nodosum ecad scorpioides in a temperate salt marsh is described. The ecad scorpioides, when transplanted from its characteristic habitat on the mid-intertidal, Spartina alterniflora-dominated, marsh flats to a location near mean low-water developed characteristics normally associated with A. nodosum ecad mackaii. The growth of these plants was more rapid than those in the mid-intertidal region and was not affected by the shading of algal fronds by S. alterniflora. Unusually high temperatures and light intensities during the winter and spring months were major factors affecting growth in plants that were subjected to relatively long periods of tidal exposure. The presence of S. alterniflora during the summer months may act in a protective capacity for mid-intertidal ecad populations.This research was supported by research grants AG-375 and BO 38018 from the National Science Foundation and, in part, by the State University of New York Research Foundation and the Energy Research and Development Administration (ERDA).     

Growth of the salt marsh periwinkleLittoraria irrorata on fungal and cordgrass diets

The growth of the salt marsh periwinkleLittoraria irrorata (collected from Sapelo Island, Georgia in 1991, initial shell length 6.2 to 11.5 mm) on various diets was measured. Growth was highest on a diet of standing-dead leaves ofSpartina alterniflora. Periwinkles provided with marsh sediment, yellow-green, sterile, or bacteria-colonized leaves lost organic mass. Fungal-colonized leaves and pure mycelia of fungi common on standing-dead leaves allowed intermediate growth. Growth onS. alterniflora-based diets was negatively correlated with the phenolics content of the food, and positively correlated with its lipid content. No correlation was found between growth and protein content. The digestibility ofS. alterniflora leaves, estimated with the acid-insoluble ash technique, was highest when yellow-green leaves were used. Colonization by fungi or bacteria caused it to decline. ForS. alterniflora-based diets, growth rates were positively correlated with the amount of time spent on the food.     

Effects of chronic oil pollution on a salt-marsh grass community

A small, naturally defined saltmarsh in Chesapeake Bay (USA) was repeatedly dosed with a No. 2 fuel oil. The production, decay rates, physical characteristics and hydrocarbon content of the dominant grass species present in the marsh were routinely monitored. Spartina alterniflora was the only grass species displaying effects of the oil dosing; this is attributed to the grass's location in the marsh. A substantial portion of the S. alterniflora in the marsh was killed by the oil dosing; the remaining portion evidenced sublethal effects including delayed development in the spring, increased density, and reduced mean weight per stem. The second annual cohort of shoots, usually produced in late summer and early fall, was suppressed almost entirely. Decay rates of S. alterniflora in the oiled marsh were higher and peaked later than decay rates in a control marsh. Oil which entered the roots and rhizomes of dead S. alterniflora was retained in a relatively undergraded state for at least 7 months after dosing stopped.Virginia Institute of Marine Science Contribution No. 945     

Grazing rates of organic matter and living fungal biomass of decaying Spartina alterniflora by three species of salt-marsh invertebrates

 The pathway for the flow of salt-marsh grass production into marsh food-webs is still not well defined. We compared the abilities of three marsh macroinvertebrates [salt marsh periwinkles, Littoraria irrorata (Say) (=Littorina irrorata), salt-marsh coffee-bean snails, Melampus bidentatus (Say); and a talitrid amphipod, Uhlorchestia spartinophila Bounsfield and Heard] to access standing-dead leaves of smooth cordgrass (Spartina alterniflora Loisel). The invertebrates were incubated with naturally-decaying leaves, and the rates of removal of organic matter and living fungal biomass (ergosterol) were measured. The impact of invertebrate activity upon fungal growth rates was measured as rates of fungal-membrane synthesis (incorporation of radioacetate into ergosterol). The removal rates of organic leaf biomass per mg individual biomass were highest for amphipods (700 μg mg⁻¹ d⁻¹) and lowest for periwinkles (90 μg mg⁻¹ d⁻¹), but the relatively large biomass of the snails made their removal rates per individual greater than those of amphipods. Net removal of ergosterol by all three invertebrates was >50% for yellow-brown (early-decay) leaf blades. For fully-brown (advanced-decay) blades, >50% removal of ergosterol was found only for periwinkles; exposure to coffee-bean snails and amphipods resulted in a net ergosterol reduction of ≤20%. The lower net reduction of living fungal biomass by coffee-bean snails and amphipods may have been due to fungal-growth stimulation (2.3-fold stimulation in coffee-bean snails and 1.5-fold stimulation in amphipods). Grazing by periwinkles did not stimulate fungal growth, possibly because of its high intensity. Grazing by these three salt-marsh shredders may affect marsh-grass shoot-decay in different ways. Periwinkles may abbreviate the period of fungal production, and incorporate the decaying material relatively quickly into snail biomass and fecal-pellet rain to the sediments. Coffee-bean snails and amphipods may enhance and prolong fungal production, along with the formation of fecal-pellet rain. All three invertebrates fed preferentially on leaf blades rather than leaf sheaths, and feeding rates of gastropods were higher during the night than during the day. Received: 25 November 1998 / Accepted: 4 November 1999     

Allocation of nitrogen and carbon in an estuarine salt marsh in Portugal

Above and below-ground biomass and nitrogen and carbon composition ofSpartina maritima, Halimione portulacoides andArthrocnemum perenne, dominating species in plant communities of the lower, middle and higher salt marsh, respectively, were compared in an estuarine salt marsh in Portugal. Plant and soil nitrogen and carbon pools were estimated. For all three species root biomass was significantly higher (70–92% of total biomass) than above-ground biomass. The percentage of root biomass was related to the location of the plants in the marsh: higher values were found in plants growing in the lower salt marsh where the sediment was more unstable and subject to tidal action, which stresses the role of the roots as an anchor. For all three species nitrogen concentrations were highest in leaves, reflecting the photosynthetic role of the tissue. For carbon higher concentrations were found in the stems, with the exception ofS. maritima. In general, lower nitrogen concentrations were found in summer, which can be explained by dilution processes due to plant growth. For both nitrogen and carbon, higher concentrations were found in the soil surface layers. Higher soil nitrogen and carbon levels were associated with higher organic matter contents. Most of the nitrogen in the salt marsh occurred in the sediments (0–40 cm) and only ca. 5.7–13.3% of the total was found in the plants. The greater portion (76.5%–86%) of carbon was found in the sediment.     

Stress tolerance of photosynthesis in sporelings of the red alga Grateloupia doryphora compared to that of Stage III thalli

More experimental evidence is needed to understand the role of propagules in macroalgal biology. There are no reports in the literature on the comparative physiology (e.g. photosynthesis) of sporelings and adults. In this paper we report on the␣variation␣in␣photosynthetic parameters (maximum photosynthesis, P _max , and efficiency, alpha, and dark respiration (R _d ) of cultivated young sporelings of the red alga␣Grateloupia␣doryphora (Montagne) Howe under normal conditions and after a short-term incubation at different salinities and temperatures. The results are compared to those␣for␣adult Stage III thalli obtained in laboratory culture from the same population of sporelings. The pigment composition of sporelings (more chlorophyll a and less phycoerthryn and phycocyanin than adults) promotes a better photosynthetic performance (higher P _max and alpha and lower R _d ) under chlorophyll a excitation. The younger sporelings were also more tolerant to variations in salinity and temperature than Stage III, in which the highest variation in maximum photosynthesis and dark respiration was observed. Received: 21 October 1996 / Accepted: 5 February 1997     

Analyzing and quantitatively evaluating the organic matter source at different ecologic zones of tidal salt marsh, North Jiangsu Province, China

In order to explore the effect of different ecological zones and their above plants in the organic matter cycling of the whole tidal salt marsh, indicators such as total organic carbon (TOC), total nitrogen (TN), C/N ratio, δ¹³C and δ¹⁵N of surface, core sediments, and plants of tidal salt marshes in North Jiangsu Province are analyzed. Subsequently, distribution regularities of these measurement indicators are discussed, and the biogeochemistry processes between sediments and plants are also analyzed. Lastly, the organic matter sources of different ecologic zones in tidal salt marsh are evaluated, and the organic matter accumulations in different ecologic zones induced by their plants are also compared. These results indicate that TOC, TN, C/N ratio and δ¹³C showed obvious zonal distribution. The organic matter sources are dominated by marine input in the silt flat, artemisia schrenkiana flat, and the transition zone between silt and spartina alterniflora flat, and are controlled by terrigenous input in spartina alterniflora flat. Spartina alterniflora plays an important role in the accumulation of organic matter in the whole tidal salt marshes ecosystem. In the study area, the annually increased TOC, organic matter and TN in the spartina alterniflora, artemisia schrenkiana and reed flats reach 6,451, 12,043 and 536 t, respectively. The amount of TOC, organic matter and TN accumulated in the spartina alterniflora flat is more than that in other ecological zones, which shows that the spartina alterniflora flat exert a non-replaceable effect on the material cycle and exchange in the whole tidal salt marshes ecosystem.     

Effects of Invasive Cordgrass on Presence of Marsh Grassbird in an Area where It Is Not Native

The threatened Marsh Grassbird (Locustella pryeri) first appeared in the salt marsh in east China after the salt marsh was invaded by cordgrass (Spartina alterniflora), a non‐native invasive species. To understand the dependence of non‐native Marsh Grassbird on the non‐native cordgrass, we quantified habitat use, food source, and reproductive success of the Marsh Grassbird at the Chongming Dongtan (CMDT) salt marsh. In the breeding season, we used point counts and radio‐tracking to determine habitat use by Marsh Grassbirds. We analyzed basal food sources of the Marsh Grassbirds by comparing the δ¹³C isotope signatures of feather and fecal samples of birds with those of local plants. We monitored the nests through the breeding season and determined the breeding success of the Marsh Grassbirds at CMDT. Density of Marsh Grassbirds was higher where cordgrass occurred than in areas of native reed (Phragmites australis) monoculture. The breeding territory of the Marsh Grassbird was composed mainly of cordgrass stands, and nests were built exclusively against cordgrass stems. Cordgrass was the major primary producer at the base of the Marsh Grassbird food chain. Breeding success of the Marsh Grassbird at CMDT was similar to breeding success within its native range. Our results suggest non‐native cordgrass provides essential habitat and food for breeding Marsh Grassbirds at CMDT and that the increase in Marsh Grassbird abundance may reflect the rapid spread of cordgrass in the coastal regions of east China. Our study provides an example of how a primary invader (i.e., cordgrass) can alter an ecosystem and thus facilitate colonization by a second non‐native species. Efectos de Spartina alterniflora Invasora Sobre Locustella pryeri en un Área Donde No Es Nativa     

Effects of tidally mediated litter moisture content on decomposition of Spartina alterniflora and S. patens

Relationships between flooding frequency, flooding duration, litter moisture levels, and litter decay rates were investigated across the natural hydrologic gradient common to intertidal salt marshes. The effects on litter decay of natural and experimental alterations of litter moisture content were assessed in both field litterbag experiments (conducted in a southern New Jersey salt marsh from 1989 to 1990) and laboratory incubations (1990). Overall, tidally mediated litter moisture content was the dominant factor controlling litter decay throughout the vegetated marsh. Rates of carbon loss were most closely related to litter moisture levels (r=0.84), which were directly related to flooding frequency (r=0.66) and duration (r=0.63). Litter moisture levels were related to elevation within the tidal range due to increasing surface levation from creekbank to high marsh (ca. 54 cm) and height of litter above the sediment surface. Carbon losses from litter of short and tall form Spartina alterniflora Loisel. and S. patens (Aiton) Muhl. along the marsh elevation gradient indicate that while some of the variations in decay rates may be due to litter type, litter moisture accounted for most of the observed variation between marsh zones and within each litter type. Mousture levels are also affected by the water retention capacity of each litter type, which may also secondarily influence decay rates. Short-term incubations of litter indicated that CO₂ evolution was positively related to moisture content with negligible respiration at moisture levels below 15% (fresh mass), increasing to a maximum between 65 and 75% depending upon litter type. Since most Spartina spp. litter remains above the marsh surface where it maintains a lower moisture content than surface litter, the use of surface litterbags may overestimate rates of carbon loss in some systems. In addition, since changes in elevation of only a few centimeters had significant effects on both litter moisture levels and decay rates, slight changes in tidal regime may have important consequences for organic matter cycling in salt marshes by affecting litter decomposition processes.     

Acetylene reduction (nitrogen fixation) in a Delaware,USA salt marsh

Acetylene reduction (nitrogen fixation) was measured in several vegetational areas in a Delaware, USA salt marsh. Samples were collected for 1 yr and the results showed a seasonally variable pattern of acetylene reduction at all stations. Peak rates were generally recorded during the later summer and early fall (September–October). The seasonality was influenced mainly, although not exclusively, by the soil temperature. In addition, samples collected in short Spartina alterniflora stands exhibited rates which were up to 20-fold higher than those found in samples from tall S. alterniflora stands. Over 50% of the total yearly ethylene production occurred from mid-August until the beginning of December at the tall and short S. alterniflora stations. Maximum activity occurred at 5 cm depth for all stations. Surface activity accounted for only 3–4% of the total measured in the top 20 cm. Addition of glucose or mannitol resulted in considerable increases in activity, thus suggesting that heterotrophic acetylene reduction is carbon and/or energy limited. The results obtained in this study indicate that the measured rates are only potential rates and that considerable caution must be used in extrapolating from acetylene reduction rates to nitrogen fixation rates in situ.     

Frog call intensities and sound propagation in the South American temperate forest region

Sound pressure levels and the spectral structure of the advertisement calls of five species of frogs from the South American temperate austral forest were analyzed. Males of Eupsophus emiliopugini, Batrachyla antartandica and B. leptopus call from the ground in bogs, while males of Hylorina sylvatica and Pleurodema thaul call from the water surface in marshes. Calling males of the species from bogs and marshes spaced at average distances that were shorter and longer than 2 m, respectively. The properties of these habitats for sound propagation were evaluated by broadcasting pure tones, broadband noise and tape-recorded advertisement calls of the three species from bogs and of H. sylvatica. Excess attenuation and spectral degradation were higher for calls broadcast in bogs than in the marsh. The calls of B.␣antartandica and B. leptopus, with dominant frequencies of about 2 kHz, were more affected than those of E.␣emiliopugini and H. sylvatica, with dominant frequencies below 1.5 kHz. These results show the lack of an optimal relationship between properties of habitats for sound transmission and the spectral structure of these anuran calls. Body size imposes an important constraint on call spectra and propagation, which frogs counteract by distribution patterns and auditory capabilities. Received: 18 April 1997 / Accepted after revision: 15 February 1998     

Flax Pond ecosystem study: Exchanges of phosphorus between a salt marsh and the coastal waters of Long Island Sound

The exchanges of phosphorus between the Flax Pond, a tidal Spartina alterniflora marsh on the north shore of Long Island (USA) and Long Island Sound were measured over 18 months. Phosphate was exported from the marsh from May through December and imported during the remainder of the year. Organic phosphorus appears to be accumulated in all seasons, but the yearly phosphorus budget of the marsh is approximately balanced despite the accumulation of about 6 mm of sediment annually.     

Food resources of postlarval brown shrimp (Penaeus aztecus) in a Texas salt marsh

Field and laboratory experiments were conducted to identify the sources of food in the natural diet of postlarval brown shrimp (Penaeus aztecus Ives). A series of enclosures placed in East Lagoon (29°20N; 94°45W) on Galveston Island, Texas, USA, in May 1985, were used to evaluate the individual and combined contribution of Spartina alterniflora detritus, epiphytes of S. alterniflora, plankton, and demersal fauna in terms of differences in shrimp growth and carbon assimilation (stable carbon-isotope analysis). Demersal fauna (harpacticoid copepods, amphipods, tanaids and polychaete annelids), and plankton (>0.095 mm) accounted for approximately 53 and 47% of the growth of the postlarvae (11 to 22 mm rostrum-telson length), respectively, while the autochthonous plant substrates, S. alterniflora detritus and epiphytes, contributed little. Laboratory experiments confirm that a mixed diet consisting of both animal protein and phytoplankton promotes maximum growth. Our results indicate that plankton may be an important allochthonous source of carbon contributing to the growth and development of shrimp in the salt marsh.     

Low molecular-weight factor from Plesiastrea versipora (Scleractinia) that modifies release and glycerol metabolism of isolated symbiotic algae

When symbiotic dinoflagellate algae (Symbiodinium sp., isolated from the coral Plesiastrea versipora) were incubated with NaH¹⁴CO₃ in the light in seawater, they released 22.69±9.16 nmol carbon/10⁶ algae. Release of photosynthetically fixed carbon was stimulated more than six-fold for algae incubated in host-tissue homogenate (148.54±97.03 nmol C/10⁶ algae) and more than four-fold (102.00±49.16 nmol C/10⁶ algae) for algae incubated in a low molecular weight fraction (≤1 000 M _r ) prepared from host homogenate. Soluble released ¹⁴C-labelled products, as determined by chromatography and autoradiography, were the same when algae were incubated in either host homogenate or the low molecular weight fraction. After 4 h incubation in the light (300 mol photons m⁻² s⁻¹),␣intracellular␣glycerol increased in algae incubated with the low molecular weight fraction (an increase of 0.39 to␣0.67 nmol glycerol/10⁶ algae) compared with little or no increase in algae incubated in seawater (0 to 0.12 nmol glycerol/10⁶ algae). Partial inhibition of triglyceride synthesis (up to 51%) was also observed when algae were incubated in the low molecular weight fraction. All these effects are the same as those observed when algae were incubated in host homogenate. These data indicate that the “host release-factor” activity of P.␣versipora is a compound of low molecular weight. Received: 13 February 1997 / Accepted: 24 October 1997     

Intertidal distribution,population dynamics and production of the amphipod Uhlorchestia spartinophila in a Georgia,USA, salt marsh

The talitrid amphipod Uhlorchestia spartinophila Bousfield and Heard occurs in close association with the smooth cordgrass Spartina alterniflora Loisel in salt marshes along the U.S. Atlantic coast. In order to assess its potential as a prey resource for secondary consumers, we followed the population and production dynamics of this amphipod from 3 November 1990 to 2 October 1991 in a salt marsh on Sapelo Island, Georgia. Semimonthly samples were taken along an intertidal transect, which extended from a vegetated creekbank levee landward 140 m to a site near the upland edge of the marsh. Amphipod densities ranged from 9 to 826 indm^-2 and were greatest in the levee and high marsh habitats at opposite ends of the transect. The highest densities occurred in March to May and the lowest in November and December. Specific growth rates, which ranged from 0.001 to 0.024 mm mm^-1 d^-1 length and 0.003 to 0.068 mg mg^-1 d^-1 AFDW (ash-free dry weight), were least in winter and decreased with increasing amphipod size. Although some reproduction occurred year-round, most of the population's reproductive output was from January to May, when adults had the greatest size-specific mass. Sex ratio usually favored females, which were larger than males. Minimum adult female size was 5.4 mm total length (TL). Broods included from 1 to 28 eggs or young and increased with increasing body size, averaging 7.4 young for an average-size female of 7.6 mm TL. Annual production, which ranged from 0.769 to 1.444 g AFDW m^-2, was least in the low marsh and greatest in the levee habitat. Production:biomass ratios were 15.4 to 17.3 in different habitats. Such high turnover rates, together with the population's broad intertidal distribution, suggest a greater contribution to trophic dynamics than is implied by the relatively low standing stock biomass of U. spartinophila in this marsh system.     

Non-indigenous ascidians in southern California harbors and marinas

Southern California's many large harbors form an important coastal ecosystem, yet they are also a␣major destination for thousands of pleasure craft and cargo vessels that have often traveled great distances. Many groups of marine organisms, including ascidians, have now been documented as undergoing range extensions as a consequence of rapid ship-transport between distant harbors phenomenon. This has resulted in a rapid increase in the rate of introductions of non-indigenous species worldwide, yet these effects of boat traffic remain largely unstudied in southern California. Ascidians are sessile marine filter-feeders, hermaphroditic, and often self-fertilizing; many species are tolerant of a wide range of environmental conditions, can reach sexual maturity in just a few weeks, and have a long breeding season. This paper documents the arrival of 14␣non-indigenous species in southern California harbors␣during this century, 13 of which have persisted:␣four prior to the 1960s (Cionaintestinalis, Styelaclava, S.␣plicata, Botryllusschlosseri), another by 1972 (S.␣canopus, formerly S. partita), and 8 since 1983 [C.␣savignyi, Ascidia zara, Ascidia sp., Polyandrocarpa zorritensis, Symplegma brakenhielmi (formerly S. oceania, and S. reptans, Microcosmus squamiger, and Molgula␣manhattensis)]. We estimate the relative abundance and seasonal fluctuations of both non-indigenous and native ascidians in all harbors in southern California from San Diego to Santa Barbara based upon the historical record, our 35 yr of field notes, and our recent surveys carried out during fall 1994, spring and fall 1995, fall 1996 and spring 1997. Possible points of origin of the exotics and predictions on further U.S. Pacific coast range-extensions are included. The concomitant decline in numbers and species of native ascidians in the harbors of southern California during this century is also reviewed. Received: 4 March 1997 / Accepted: 26 September 1997     

Effects of ambient ammonia levels on blood ammonia, ammonia excretion and heart and scaphognathite rates of Nephrops norvegicus

During commercial handling of Nephropsnorvegicus (L.) there are a number of situations when the prawns may be exposed to very high ambient ammonia levels. These experiments evaluated the effects of increased levels of ambient total ammonia (TA = NH₃ + NH₄ ⁺) on␣blood ammonia, ammonia efflux rates and on the cardio-ventilatory performance of N. norvegicus. When prawns were taken from <1 to 2000 μmol TA l⁻¹ medium, blood TA concentrations increased rapidly for the first 2 h but tended to drop thereafter. Original blood TA levels were restored 6 h after the prawns were transferred back from seawater containing 2000 to <1 μmol TA l⁻¹. Sudden exposure to 500, 1000, 2000 or 4000 μmol TA l⁻¹ medium induced blood TA concentrations to increase respectively to 50, 30, 33 and 36% of external concentrations (normally, internal TA values are much higher than external levels). Immediately after transfer back to seawater with low ammonia concentration ( <1 μmol TA l⁻¹), excretion rates were higher than those of control prawns, and the absolute amounts of TA excreted were considerably higher than those calculated to have accumulated in the haemolymph. Heart rate (HR) and scaphognathite rate (SR) were not altered when prawns were subjected to sudden alterations in ambient ammonia ( <1 to 2000 to <1 μmol TA l⁻¹). When water ammonia concentrations were altered more gradually, both rates increased, but only at 4000 μmol TA l⁻¹. These results show that N. norvegicus is able to remove ammonia from the haemolymph and/or transform ammonia into some other substance when subjected to increased levels of ambient ammonia. Possible mechanisms involved (e.g. active transport across the gills; storage in some other tissue; glutamate synthe sis) are discussed. Received: 20 May 1996 / Accepted: 1 July 1996     

Habitat utilization and alteration by the invasive burrowing isopod, Sphaeroma quoyanum, in California salt marshes

In recent years the pace of exotic species introduction and invasion has accelerated, particularly in estuaries and wetlands. Species invasions may affect coastal ecosystems in many ways. Alteration of sedimentary environments, through structure formation and burrowing, has particularly dramatic effects on coastal habitats. This study examines modification of channel bank and marsh edge habitat by the burrowing Australasian isopod Sphaeroma quoyanum Milne Edwards, in created and natural salt marshes of San Diego Bay and San Francisco Bay. Abundance and distribution patterns of this isopod species, its relationships with habitat characteristics, and its effects on sediment properties and bank erosion were examined seasonally, and in several marsh microhabitats. Mean isopod densities were 1541 and 2936 individuals per 0.25 m² in San Francisco Bay, and 361 and 1153 individuals per 0.25 m² in San Diego Bay study sites during December and July 1998, respectively. This isopod forms dense, anastomosing burrow networks. S. quoyanum densities did not differ as a function of location within creeks or location in natural versus created marshes. Burrows, which are on average 6 mm wide and 2 cm long, were associated with firm sediments containing high detrital biomass. Although erosion is a natural process along salt marsh banks, enclosure experiments demonstrated that isopod activities can enhance sediment loss from banks. In areas infested with S. quoyanum, losses may exceed 100 cm of marsh edge per year. The effects of habitat alteration by this invading species are likely to increase in severity in the coastal zone as these ecosystems become degraded. Received: 30 March 2000 / Accepted: 21 September 2000