Positive effects of the introduced green alga, Codium fragile ssp. tomentosoides, on recruitment and survival of mussels

The green macroalga, Codium fragile ssp. tomentosoides, is an important component of sheltered low-shore assemblages on breakwaters along sandy shores in the northern Adriatic Sea. Macroscopic thalli of C. fragile are not perennial, but develop from propagules and/or undifferentiated forms in early spring, when the dominant native space-occupier, the mussel Mytilus galloprovincialis, recruits. By mid-summer, rapid growth of C. fragile leads to the formation of a dense canopy. We investigated the effects of juvenile and adult thalli of C. fragile on recruitment, survival and growth of mussels. Two experiments tested the hypotheses: (1) that recruitment of mussels is greater within patches of juvenile thalli (primordia) of C. fragile than on adjacent bare surfaces; (2) that the presence of a canopy of C. fragile affects the survival and growth of mussel recruits. The number of recruits of mussels was significantly larger within clumps of primordia of C. fragile than on bare surfaces. The removal of the canopy of C. fragile affected negatively the density of mussels after 2 months from the start of the experiment, but there were no effects on the mean size of individuals, nor on the size–frequency distribution. The same trend persisted after 4 months from the start of the experiment. These results show that re-colonisation of space by mussels is enhanced by C. fragile. Given that mussels, in turn, have the potential to reduce recruitment rates of C. fragile, quick recovery of mussel beds after disturbances could be crucial for controlling the abundance of this alga on breakwaters. Results also suggest that the effects of introduced species on native assemblages can be explained only through studies encompassing different life-stages of interacting organisms.     

Growth patterns of Codium fragile ssp. tomentosoides in response to temperature,irradiance, salinity,and nitrogen source

Seasonal patterns of growth, reproduction, and productivity of Codium fragile spp. tomentosoides (van Goor) Silva were monitored at 3 locations in Rhode Island. Maximal growth occurred during the summer and was more significantly correlated with temperature than any other factor measured in this study. Multiple correlation models suggested an interaction between temperature, irradiance, and available nitrogen. Maximal reproduction occurred in late summer and early fall. The maximal productivity, based on harvested quadrats, was 2. 10 g dry weight m^-2 day^-1. A large amount (up to 87.3%) of the annual production entered the detrital food chain during the winter by fragmentation of the thallus. Culture studies examined the effects of temperature (6° to 30°C), irradiance (7 to 140 E m^-2 sec^-1), daylength (8 h light: 16 h dark to 24 h light: O h dark) and salinity (6 to 48) on growth. Differentiated thalli grew over a broad range of experimental conditions, with maximal growth at 24°C, 24 to 30 S, a minimal irradiance of 28 E m^-2 sec^-1, and 16 h daylength. The effect of increasing daylength was due to increased total daily irradiance rather than to a true photoperiodic effect. Undifferentiated sporelings survived and grew in a narrower range of environmental conditions than thalli. c. fragile spp. tomentosoides grew equally well with nitrate, nitrite, ammonium, and urea as a nitrogen source. The addition of NaHCO₃ stimulated growth at levels of 2.4 to 4.8 mM, suggesting an inorganic carbon limitation in static cultures. This study supports the hypothesis that the in situ seasonal growth pattern of c. fragile spp. tomentosoides is primarily due to the interaction of temperature and irradiance.     

Comparative feeding preference of Strongylocentrotus droebachiensis (Echinoidea) for the invasive seaweed Codium fragile ssp. tomentosoides (Chlorophyceae) and four other seaweeds

Laboratory experiments conducted during 1987 on Appledore Island, Maine, USA, tested whether feeding preference or the absence of an attractant was the cause for the occurrence of beds of Codium fragile ssp. tomentosoides (herein referred to as Codium fragile) within rocky barrens grazed clear of kelp by the sea urchin Strongylocentrotus droebachiensis. Consumption of C. fragile in single-diet experiments (1 seaweed/sea urchin) was highly variable and was not significantly different from that for several other seaweeds (Agarum cribrosum, Ascophyllum nodosum, Chondrus crispus, and Laminana saccharina) important in the field diet of the green sea urchin. In multiple-diet experiments (5 seaweeds/sea urchin) significantly less Codium fragile was eaten than Chondrus crispus, but significantly more Codium fragile was eaten than A. cribrosum. Chemosensory experiments suggest that C. fragile does not attract the sea urchin. Sea urchins are unable to detect C. fragile but will eat it when they come in contact with it.     

Introduced versus native subspecies of Codium fragile: how distinctive is the invasive subspecies tomentosoides?

After its introduction, the green alga Codium fragile (Sur.) Hariot ssp. tomentosoides (van Goor) Silva has spread widely on several temperate-zone, rocky shores where non-weedy conspecific subspecies occur (N.E. Atlantic, N.E. Pacific, S. Pacific). To determine how phenologically and morphologically distinctive the invasive alga was relative to native subspecies, I compared marine intertidal populations of C. fragile ssp. tomentosoides and the native C. fragile ssp. novaezelandiae (J. Ag.) Silva (hereafter referred to as ssp. tomentosoides and ssp. novae-zelandiae respectively on New Zealand shores in 1992, 1993 and 1995. On the North Island, the invasive ssp. tomentosoides is sparsely distributed on low intertidal benches on wave-protected shores in the Hauraki Gulf (east coast) in spring and summer, and thalli die back to the perennial holdfast in autumn. In contrast, the native ssp. novaezelandiae forms dense beds within the low intertidal mussel zone on wave-swept shores of Maori Bay (west coast), and fronds are perennial. Whereas ssp. tomentosoides has only a few fronds arising from the spongy basal hold-fast, ssp. novae-zelandiae thalli are composed of many fronds. The ssp. tomentosoides from the Hauraki Gulf is significantly more branched than comparably sized native conspecifics from Maori Bay. These phenological and morphological differences were used to predict the subspecific identity of C. fragile from three other locations on the North Island, two locations on the South Island, and four locations on S.E. Australian shores; microscopic examination of utricles was used to check the predictions. Seasonality and number of fronds per thallus are the most reliable characters for field identification of native vs invasive subspecies: perennial intertidal thalli with large numbers of fronds are indicative of native subspecies for different geographic regions.     

Feeding, growth and reproduction of sea urchins (Strongylocentrotus droebachiensis) on single and mixed diets of kelp (Laminaria spp.) and the invasive alga Codium fragile ssp. tomentosoides

Since its introduction to Nova Scotia in the early 1990s, the invasive green alga Codium fragile ssp. tomentosoides has spread rapidly to become a dominant and persistent component of seaweed assemblages in the rocky subtidal zone. To examine the potential of sea urchins (Strongylocentrotus droebachiensis) to regulate Codium, and the potential of the alga to support urchin growth and reproduction, we fed urchins Codium and/or kelp (Laminaria spp., a high quality and preferred native food) in the laboratory for 11 months. Urchins showed a strong and active preference for kelp: they consumed more Laminaria than Codium (as dry weight) in single and mixed diet treatments. Urchins fed only Codium consumed 3.5 times more of the alga than those on a mixed diet, but did not increase their feeding rate in winter when kelp consumption was greatest. Laminaria was consumed at the same rate in single and mixed diets, indicating that the presence of an alternative food had no effect on kelp consumption. Survival and growth (change in test diameter) were lowest on the Codium diet, although the differences among diets were not statistically significant. Urchins on the Codium diet showed no gonadal production over the annual reproductive cycle, compared to a marked rise in gonad index on the Laminaria and mixed diets. Our results indicate that Codium is an unattractive, unpalatable and poor quality food, which is not readily consumed by urchins. Thus we predict that urchins at moderate densities will be much more likely to graze kelp than Codium in mixed stands, creating a mosaic of Codium patches and barren areas. At high densities, however, urchins are likely to destructively graze all seaweeds, although dense stands of the invasive alga may prolong the widespread formation of barrens.     

Ecological studies of Codium fragile in New England,USA

The seasonal growth and reproduction of Codium fragile were studied at Boothbay Harbor, Maine, and at Woods Hole and Wings Neck, Massachusetts (USA), in relation to several environmental parameters. Maximum growth and reproduction occur during periods of highest water temperatures and insolation. The effects of various light intensities and temperatures on the photosynthesis rates of C. fragile were determined and correlated with the field results. Optimal conditions for net photosynthesis are 21° to 24°C and 900 to 1100 foot-candles. It is suggested that C. fragile is a warm-temperate plant that is growing near its northern limits in New England, but one that has not extended to its potential southern limits.Published with the approval of the Director of the New Hampshire Agricultural Experiment Station as Scientific Contribution Number 625.     

Nitrogen fixation by blue-green algae associated with the siphonous green seaweed Codium decorticatum: effects on ammonium uptake

Nitrogen fixation (acetylene reduction) at rates of up to 1.2 g N₂ g dry wt^-1 h^-1 was measured for the siphonous green seaweed Codium decorticatum. No nitrogenase activity was detected in C. isthmocladum. The nitrogenase activity was light sensitive and was inhibited by the addition of DCMU and triphenyl tetrazolium chloride. Additions of glucose did not stimulate nitrogen fixation. Blue-green algae (Calothrix sp., Anabaena sp., and Phormidium sp.) were implicated as the organisms responsible for the nitrogenase activity. They occurred in a reduced microzone within the C. decorticatum thallus where nitrogen fixation was optimized. Nitrogen fixation did not affect the kinetic constants for ammonium uptake in C. decorticatum (K_s=12.0 M, V_max=13.4 mol NH₃ g dry wt^-1 h^-1) determined using the perturbation method. Nevertheless, C. decorticatum thalli which fixed nitrogen had internal dissolved nitrogen concentrations which were over 1.4 times higher than in non-fixing thalli. This suggests that if C. decorticatum does derive part of its nitrogen requirement from the blue-green algae which it harbors, the transfer does not involve competition between this process and the uptake of ambient ammonium.     

Comparisons in demographic rates of bay scallops in eelgrass and the introduced alga, Codium fragile, in New York

Bay scallops, Argopecten irradians, supported vibrant fisheries which subsequently collapsed, as such, they are a focus species for many restoration efforts along the Atlantic and Gulf coasts of the United States. The scallops’ preferred habitat, seagrass, has also dramatically declined, and some scallop populations have increased post-restoration despite reduced seagrass cover. This has led to the hypothesis that macroalgae may serve as suitable alternative habitats for bay scallops. This study is the first to compare demographic rates, such as long-term survival, growth, condition and reproductive potential of scallops between the native eelgrass, Zostera marina, and the introduced alga, Codium fragile. Although long-term survival was not different between habitats, results suggest site-specific and inter-annual variation in the impacts of Codium on scallop growth. While demographic rates did not differ in Shinnecock Bay, in Sag Harbor, growth and/or condition were significantly different between both vegetated habitats depending on the year. However, recruit density, size and condition did not vary significantly, adding to the complexity of this relationship. Despite potential site-specific and inter-annual differences, this study supports the hypothesis that habitats other than eelgrass can benefit bay scallops.     

Comparative survival of bay scallops in eelgrass and the introduced alga, Codium fragile, in a New York estuary

Eelgrass, Zostera marina, is generally regarded as the preferred habitat of bay scallops, but in some cases scallop populations have persisted or increased in areas lacking eelgrass. This suggests that some other substrate(s) may serve important ecological functions for bay scallops. One candidate is Codium fragile, a macroalgal species with which bay scallops are known to associate and in which we commonly find juvenile and adult bay scallops in eastern Long Island, New York. In this study, we examined whether survival of planted bay scallops differed in Codium, eelgrass, and Codium + eelgrass substrates at two sites during August and October of 2 years. Survival of tethered scallops and recoveries of live free-planted individuals varied with scallop size, planting season and year, but no differences were observed between the three substrates for a given scallop size and planting date. Crab (particularly Dyspanopeus sayi) and whelk predation were implicated as important causes of tethered scallop mortalities while emigration and removal by predators likely contributed to scallop losses. Densities of naturally recruited 0+ years scallops recovered by visual and suction dredge sampling were similar in the eelgrass and Codium substrates. While our results suggest that Codium may offer some degree of predation refuge for bay scallops, further work needs to weigh the potential disadvantages of this substrate (such as low DO levels, potential attachment and transport of scallops, and differences in current flow, food availability and sedimentation relative to eelgrass) to determine if Codium may serve as a valuable habitat for bay scallops throughout their lifespan.     

Homeward paths of displaced juvenile alligators as determined by radiotelemetry

Summary To test their ability to home without searching or wandering, 19 juvenile alligators were radiotracked after displacement to areas 1–10 home range diameters beyond their areas of familiarity. Movement rates varied inversely with vegetation density, and in some cases habitat barriers appeared to deflect an alligator's homeward path. However, failures to home occurred only in those cases where major habitat barriers separated an alligator from its home site. In all other cases the alligators headed directly homeward; at least 10 of these completed their homeward journeys. The orienting ability of the alligators was neither dependent on the conditions under which they were displaced, nor the environmental conditions prevailing after release. These observations support the hypothesis that alligators possess a true navigational ability.     

Nitrogen limitation of net primary productivity in terrestrial ecosystems is globally distributed

Our meta-analysis of 126 nitrogen addition experiments evaluated nitrogen (N) limitation of net primary production (NPP) in terrestrial ecosystems. We tested the hypothesis that N limitation is widespread among biomes and influenced by geography and climate. We used the response ratio (R approximately equal ANPP(N)/ANPP(ctrl)) of aboveground plant growth in fertilized to control plots and found that most ecosystems are nitrogen limited with an average 29% growth response to nitrogen (i.e., R = 1.29). The response ratio was significant within temperate forests (R = 1.19), tropical forests (R = 1.60), temperate grasslands (R = 1.53), tropical grasslands (R = 1.26), wetlands (R = 1.16), and tundra (R = 1.35), but not deserts. Eight tropical forest studies had been conducted on very young volcanic soils in Hawaii, and this subgroup was strongly N limited (R = 2.13), which resulted in a negative correlation between forest R and latitude. The degree of N limitation in the remainder of the tropical forest studies (R = 1.20) was comparable to that of temperate forests, and when the young Hawaiian subgroup was excluded, forest R did not vary with latitude. Grassland response increased with latitude, but was independent of temperature and precipitation. These results suggest that the global N and C cycles interact strongly and that geography can mediate ecosystem response to N within certain biome types.     

Resource partitioning by reef corals as determined from stable isotope composition

The pattern of resource partitioning vs depth by corals collected in February 1983 from Jamaica and the Red Sea was determined from their stable carbon isotope composition. Observations were made on isolated zooxanthellae and corresponding algae-free animal tissue from eight species at four depths over a 50 m bathymetric range. Zooxanthellae ¹³C was high in shallow water and became lower as depth increased. This trend correlated significantly with the anual integrated photosynthetic rate. The trend is interpreted according to a depletion-diffusion hypothesis; in shallow water, at high rates of photosynthesis, metabolic CO₂ is nearly depleted and the supply of CO₂ from seawater bicarbonate is limited by diffusion. Since most of the available CO₂ is fixed, isotope fractionation is minimal. In deeper water, at lower rates of photosynthesis, metabolic CO₂ is ample, and isotope fractionation is greater. Animal tissue ¹³C was slightly lower than corresponding zooxanthellae values in shallow water. As depth increased the difference between zooxanthellae and animal tissue ¹³C increased and the latter approached the ¹³C of oceanic particulate organic carbon. These data suggest that carbon is translocated at all depths and that deep-water corals draw significantly on allocthonous sources of carbon.Contribution No. 436 of the Discovery Bay Marine Laboratory of the University of the West Indies     

Trophic diversity in amphipods within a temperate eelgrass ecosystem as determined by gut contents and C and N isotope analysis

A conjoint analysis of gut contents and stable C and N isotopes was applied to determine the main food sources and feeding habits of dominant amphipods in an eelgrass bed (Zostera marina) in Gwangyang Bay, Korea. Gut content observations demonstrated that, while Gammaropsis japonicus and Jassa slatteryi are herbivorous, feeding on epiphytes and detritus, Pontogeneia rostrata and Monocorophium acherusicum are omnivorous, feeding on mesozooplankton fragments and detritus. Stable isotope data confirmed that epiphytes, detritus, and mesozooplankton fragments were major food sources for amphipods in the eelgrass bed. Isotopic mixing model calculations clearly showed an interspecific difference in diet composition. A high isotopic dissimilarity between amphipod taxa demonstrated interspecific trophic diversity, reflecting their herbivorous (G. japonicus and J. slatteryi) and omnivorous (P. rostrata and M. acherusicum) feeding habits and confirmed the detrivorous feeding habits of caprellids. Such trophic diversity at interspecific level of the amphipod species indicates that they use different food resources within their microhabitats and play species-specific functional roles as mediators in trophic pathways from producers to higher-level consumers of the eelgrass ecosystem. Finally, our findings suggest that information on the species-specific trophic ecology of amphipods is needed to better understand their potential role in the trophic dynamics and carbon flow of seagrass bed ecosystems.     

不同施氮水平条件下烤烟对氮素的利用研究

大田中设置盆栽试验，采用１５Ｎ示踪法对不同施氮水平条件下的烤烟的氮素吸收、肥料氮素的利用和土壤ＡＮ值进行研究．试验结果表明，烤烟全生育期吸收的氮素主要来自于土壤可利用性氮．随着氮肥用量的增加，烤烟对肥料氮的吸收量增加，对土壤氮的吸收量减少，土壤对肥料氮素的固定和氮肥损失量增加．ＡＮ值有随氮素用量的增加而逐渐减小的趋势，但以处理４的（６７．５ｋｇ氮素／ｈｍ２）最小．肥料中的氮素利用率以处理４最高，达到５２．３％。     

Russell's definite description analysis of production and limitation of phytoplankton species

The majority of phytoplankton species observed in the southwestern Atlantic Ocean produce new cells to equal cells lost, if any, to grazing and sinking. At the same time these species must limit new cells to numbers equalling any lost, since their abundance remains about the same in both deep and shallow areas. Of each of these species one may assert: the producer is a limiter (1). One species, coccolithus huxleyi, produces cells to offset any lost in deep water but does not limit, as it drifts into shallow water, new cells to equal the number lost in deep water, since it is more abundant in shallow than in deep water. For C. huxleyi it is true that the producer is not a limiter (2). However, once in the shallows, C. huxleyi is in a steady state, so that nutrient concentration both produces and limits new cells to equal any lost. Thus, (2) converts to (1) by having nutrient concentration the producer and the limiter. “The producer is a limiter” is a covering statement. Asserting it covers all observable species; denying it covers species not observed, those rare species which can only be found by more exhaustive sampling.     

Feeding habits of the dungeness crab Cancer magister as determined by the index of relative importance

Specimens of the Dungeness crab Cancer magister, were collected subtidally and intertidally from an estuary in Washington State, USA in June and September 1980, and January, April, May and July, 1981. Gut contens of freshly collected crabs were analyzed by the Index of Relative Importance; for each prey taxon, this method measured frequency of occurrence, percentage of total biomass, and percentage of total numbers consumed. The most important higher taxon eaten was fish; however, the most important prey genus was the shrimp Crangon spp. There was greater predation on Crangon spp. at night at the intertidal site, and during winter and spring when the shrimp were most abundant there. Feeding activity, as indicated by a weight-specific gut-fullness index, showed no consistent diel pattern. There were significant ontogenetic changes in feeding patterns: first-year crabs preyed primarily on very small bivalves or small crustaceans including their conspecifics; second-year crabs preferred Crangon spp. and fish, and third-year crabs preyed less on Crangon spp. and more on fish. Such changes in feeding habits with ege could be purely due to mechanics of food handling, but might also reduce competition among age groups of crabs, possibly partitioning resources within the estuary. Findings are discussed in terms of optimal foraging and compared to other similar studies.Contribution No. 599, School of Fisheries, University of Washington, Seattle, Washington 98195, USA     

一株假单胞菌WSH1001的脱氮特性

研究了假单胞菌WSH 1001(Pseudomonas sp.WSH 1001)对氨氮及硝态氮去除性能的影响因素以及WSH 1001在实际污水处理中的应用情况,并将其与市售硝化菌制剂的脱氮性能进行了比较.结果表明:菌株前培养方式对后续的氮去除性能影响较大,葡萄糖或柠檬酸钠是最适碳源;在20～35℃的范围内,温度对氨氮及硝态氮的去除率没有明显影响;溶氧浓度对氨氮及硝态氮的去除效率影响很大;金属离子Cu2+、Co2+和Zn2+极大地抑制了该菌株对氨氮及硝态氮的去除能力;菌株WSH 1001在6 h内对70 mg L-1的氨氮去除率高达99.64%,总氮去除率达94.94%,在8 h内对50 mgL-1的硝态氮去除率达到了87.69%,说明该菌株同时具备硝化和反硝化的能力;当菌株WSH 1001应用于实际污水(初始氨氮和COD浓度分别为44和113 mg L-1)、并额外添加3 g L-1的丁二酸钠作为外加碳源时,氨氮去除率在6 h时达到99.23%,较其它市售硝化菌制剂脱氮性能高.该研究表明假单胞菌WSH 1001在实际污水的处理上具有较好的应用潜能.     

Dispersal potential of invasive algae: the determinants of buoyancy in <Emphasis Type="Italic">Codium fragile</Emphasis> ssp. <Emphasis Type="Italic">fragile</Emphasis>

The capacity for long-distance dispersal is an important factor in determining the spread of invasive species. For algae, positive buoyancy generally is correlated with increased dispersal potential, and the light environment has been previously identified as a possible determinant of buoyancy in several species. We examined the effect of light intensity on the buoyancy of fragments of the invasive green alga Codium fragile ssp. fragile. Under natural and controlled conditions, the buoyancy of samples taken from the thallus tip was higher than those from near the holdfast. Both laboratory and field experiments also showed that buoyancy was dynamic and switched from positive to negative under reduced light intensity, but this change required several days. We also observed seasonal changes in buoyancy, presumably due to natural variations in light intensity, with the buoyancy of fragments washed up on the shore highest in mid-summer. These results show that buoyancy is a dynamic property of the C. fragile ssp. fragile thallus and suggest that buoyant fragments contribute to long-range dispersal and accelerated regional spread of this invader. This finding suggests that dispersal is more likely during conditions of high light intensity and illustrates the need to understand how variations in the natural environment can affect the dispersal potential of invasive species.