Incorporation of organic aggregates by marine mussels

Two marine mussels, Geukensia demissa (Dillwyn) and Mytilus edulis (L.) collected in 1990 in Old Silver Beach, Falmouth, Massachusetts, incorporated nitrogen when fed ¹⁵N-labelled organic aggregates produced from dissolved organic nitrogen released by the brown sea-weed Fucus vesiculosis. Uptake of ¹⁵N on the aggregate diet was linear over the course of 24 h, and unincorporated ¹⁵N was eliminated from the gut after 48 h. Both species of mussels incorporated approximately five times more N when they were fed organic aggregates than when they were fed either ¹⁵N-labelled dissolved organic material (DOM) or particulate detritus, both of which were also derived from the seaweed. Nitrogen uptake was greatest in controls fed the diatom Thalassiosira weissflogii; mussels fed phytoplankton incorporated seven times more nitrogen than those fed aggregates. However, aggregates could supply an estimated 7 to 14% of the N requirements for both mussels, whereas DOM or particulate detritus could only supply 1 to 3%. These data provide evidence that a food web pathway exists from primary producer to released dissolved organic nitrogen to microbial organic aggregate to metazoan consumer, and, further, that it can be more important in a detrital food web than either particulate detritus or DOM.     

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.     

Effect of food type and ration on growth of juvenile Capitella sp. I (Annelida: Polychaeta): macro- and micronutrients

Growth rates of juvenile Capitella sp. I were determined on different rations of six food types: Gerber's mixed cereal, TetraMin fish flakes, benthic diatoms, Ulva sp., spring detritus, and summer detritus. A simple growth model estimated maximum growth rate and maintenance ration for each food. There were differences in the growth response among foods relative to nitrogen content. As juveniles increased in size, differences in growth between foods became more pronounced. For all juveniles, growth rates were correlated with levels of the amino acids histidine, phenylalanine, threonine, and valine, and the polyunsaturated fatty acid 20:5w3; correlations with histidine and phenylalanine levels were the most significant. Regressions of growth rates as a function of these two amino acids suggest a daily maintenance ration of 300 pg histidine and phenylalanine mg^-1 nitrogen biomass. Juvenile worms grew on spring detritus but not on summer detritus, indicating the probable importance of micronutrients (polyunsaturated fatty acids, amino acids) for growth of juvenile Capitella sp. I in the field.     

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).     

Relative importance of benthic microalgae,phytoplankton, and mangroves as sources of nutrition for penaeid prawns and other coastal invertebrates from Malaysia

Carbon, sulfur, and nitrogen stable isotope ratio techniques were used in 1989 and 1990 to evaluate the relative importance of algae and of mangrove detritus in the nutrition of two penaeid prawn species on the west coast of Peninsular Malaysia. Mangrove detritus was found to contribute to the nutrition of juvenile Penaeus merguiensis de Man living within tidal creeks, but not to adult P. merguiensis and juvenile and adult Parapenaeopsis sculptilis (Heller) captured offshore. Results from radiotracer feeding studies, with refractory ¹⁴C mangrove lignocellulose as the food source, indicated that juvenile P. merguiensis from tidal creeks assimilated mangrove carbon with an efficiency of 13.4%. This did not differ significantly (P=0.05) from the assimilation efficiencies of juvenile and adult P. sculptilis living offshore (10.0 and 10.9%, respectively); these values were significantly higher (P<0.001) than for adult P. merguiensis (2.1%). Thus, the differential contribution of mangrove material to the nutrition of prawns in the tidal creeks and offshore was not related to differences in the prawn's ability to utilize detritus. Instead, our analysis of C stable isotopes in sediments indicated that mangrove detritus was generally more abundant within the tidal creeks than offshore. Juvenile prawns in the tidal creeks may also utilize mangrove material indirectly by feeding on small detritivorous invertebrates. Stable isotope analyses suggest that benthic microalgae constitute the other major dietary component for prawns living in tidal creeks. Prawns offshore were utilizing mainly phytoplankton-based material.     

Growth,food utilization and effect of social interaction in the European bass Dicentrarchus labrax

Fingerling and juvenile bass, Dicentrarchus labrax (L.) fed on filleted saithe, Gadus virens, exhibited much higher K ₁ energy and nitrogen utilization efficiencies (growth as % energy or nitrogen intake) than those fed on trout pellets or shore crabs, Carcinus maenas, although the efficiency declined during prolonged feeding. Mean organic absorption efficiency for saithe was 99.5% and was independent of food intake but the efficiency for pellets and crab declined with increasing food intake from 99 to 92%. For each diet the regression of relative growth rate on food intake was linear up to the regression of relative growth rate on food intake was linear up to the highest feeding level and independent of body weight, enabling construction of a simple model for food transformation. The regression coefficient (equivalent to K ₃, the ratio of growth to food consumed in excess of maintenance) for saithe was about 3 times that for pellets in terms of energy and twice that for pellets in terms of nitrogen. The more efficient utilization of saithe for growth and of pellets for maintenance were related to differences in their nutritional value. With each diet, K ₃ and maximum K ₁ for energy were about twice those for nitrogen. Social interaction had a significant effect on food utilization in juvenile bass. Isolated fish showed poor growth and mean K ₁ efficiency in comparison with bass in pairs or groups. K ₃ efficiency and mainternance requirement were similar in individuals and groups, but were significantly greater in pairs. These results are related to behavioural observeations and environmental stress, and the limitations of extrapolating the results of laboratory feeding experiments on fish to other situations is discussed.     

Use of an inert radioactive particle for measuring particle accumulation by filter-feeding bivalve molluscs

Because the rate of microbial degradation differs for the various sources contributing to the detrital pool in marine systems, their availability to detritivores might also vary. Carbon-14 tracer experiments were used to compare differences in the oxidation and net incorporation by the polychaete Capitella capitata of a nitrogenrich, easily-decomposable detritus derived from the red macrophytic algae Gracilaria sp. versus a nitrogen-poor, decay-resistant detritus derived from the eelgrass Zostera marina. The net incorporation of Gracilaria sp. detritus by C. capitata reached a maximum (91 g dry weight of detritus/mg dry weight of worm/day) after only 14 days of decomposition, whereas that of z. marina detritus equaled this level after 30 days of aging, but continued to increase to 375 g at 180 days. The oxidation rate of Gracilaria sp. detritus was consistently higher (peak of 61 mg dry weight of detritus/day at 30 day-aging) than z. marina detritus, which reached this level only after 180 days of aging. The presence or absence of C. capitata did not significantly alter the oxidation rate. The above difference might be attributed to a rapid exploitation and mineralization by bacteria of the more available Gracilaria sp. detritus, but a slow, controlled utilization by the microbes of the less available Z. marina detritus, especially during the early stage of decomposition. This would allow maximum exploitation of the substrate by macroconsumers, resulting in more of the detrital resource being tied up in detritivore biomass rather than being rapidly mineralized to CO₂. Difference in the length of aging at which various detrital sources become available to detritivores could result in a temporal partitioning of food resources and should be considered in attempting to understand the dynamics of detrital-based food chains.     

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.     

Herbivory and detritivory among gammaridean amphipods from a Florida seagrass community

The gammaridean amphipods Cymadusa compta (Smith), Gammarus mucronatus Say, Melita nitida Smith and Grandidierella bonnieroides Stephensen from a seagrass community in the Indian River estuary of Florida (USA) fed variously upon large drift algae, small algae epiphytic on seagrasses and seagrass leaf debris and detritus. Consumption was measured in the laboratory using an index (CI) equivalent to mg ingested mg^-1 amphipod day^-1. Observations revealed that the amphipods fed by macrophagy, an attack upon large algae and seagrass debris, and by microphagy, small particle detritus feeding and scraping of plant surfaces for diatoms and other epiphytic algae. C. compta was a macrophagous feeder with a generalized diet of algae and seagrass debris, preferring epiphytic algae and drift algae at mean rates of 1.10 and 0.87 CI, respectively. Gammarus mucronatus fed upon epiphytes and seagrass debris equally at mean rates of 0.90 and 0.97 CI, respectively. The diet of M. nitida condisted primarily of epiphytes, consumed at an average rate of 1.05 CI. Grandidierella bonnieroides fed in a specialized microphagous manner, grooming plant surfaces for small particle detritus and diatoms at an approximate CI rate of 1.45. Assimilation of plants ingested, as reflected by carbon-14 uptake, varied similarly among the 4 amphipods. Epiphytic algae appeared to be most useful as food, providing means of 41 to 75% carbon-14 uptake as ingesta. Drift algae and seagrass debris were of less value, with means varying between 11 and 24 % of carbon-14 uptake by the amphipods. The data show a pattern of feeding which resembles resource partitioning of food both by size and kind. Other evidence, however, including population limitation by predators and an apparent overabundance of food, indicate that resource partitioning as seen may be an artifact, and one which has no co-evolutionary basis among the present species.Contribution No. 102 of Harbor Branch Foundation, Inc.     

An assessment of seaweed decomposition within a southern Strait of Georgia seaweed community

An assessment of litter and detritus decomposition and nitrogen content of decomposing litter is presented for ten important seaweeds within a southern Strait of Georgia (British Columbia, Canada) seaweed community sampled from August 1975 until October 1976. Litter decomposition rates varied among species with the time required for litter to disappear from litter bags ranging from 6 d for the lamina of Nereocystis luetkeana to about 70 d for Fucus distichus. Decomposition was characterized by an accelerating increase in the nitrogen: dry weight ratio of remnant litter as decomposition proceeded. Iridaea cordata detritus decomposed most rapidly, at 5.7% d^-1, while rates for Gigartina papillata, N. luetkeana, Laminaria saccharina and Laminaria groenlandica ranged from 1.8 to 3.6% d^-1. The remaining species decomposed more slowly. There was a tendency toward more rapid decomposition with decreasing crude fibre content and detritus particle size; however, it appears that morphology, habitat and growth rate are also correlated with relative decomposition rates. Of 43 taxa identified within quantitative litter collections, F. distichus (41%), I. cordata (26%), N. luetkeana (27%) and Laminaria spp. (4%) accounted for 98% of total deposition with mean peak accumulation occurring in August and September from a low near zero in January and February. Litter distribution was patchy, with most litter decomposing near its place of deposition. The application of litter decomposition rates to measured litter accumulation in a mathematical simulation of decomposition predicted the rate of seaweed litter decomposition to peak at about 1.1 g AFDW (ash-free dry weight) m^-2 d^-1 in early September from a mid-winter low near zero. In total, 56±4% of decomposing litter formed detritus, with the remainder being released as soluble matter. The annual contribution of seaweed litter biomass to detrital pathways from our study site was calculated to be 152 g AFDW m^-2.     

Survival, growth and reproduction of the salt-marsh amphipod Uhlorchestia spartinophila reared on natural diets of senescent and dead Spartina alterniflora leaves

The talitrid amphipod Uhlorchestia spartinophila lives in close association with standing-dead leaves of the smooth cordgrass Spartina alterniflora Loisel in salt marshes along the Atlantic coast of North America. This study probed the strength of the trophic link between the amphipod population and the decomposition process in this detrital-based ecosystem. We measured survival, growth and reproductive output in groups of amphipods reared for 6 wk on five diets derived from sheath and blade portions of S. alterniflora leaves just prior to (senescent) and during (dead) decomposition. In unfed treatments, the daily specific mortality rate was 0.391 and starved amphipods survived no longer than 11 d. Among the fed treatments, a diet of senescent sheaths resulted in the lowest survival (20%) and yielded no offspring. Groups fed senescent blades, dead sheaths, dead blades and unwashed dead sheaths had survival rates of 56 to 84% and produced 5.0 to 12.5 offspring replicate⁻¹. Sex ratio usually favored females, but approached unity in treatments with high overall survival, suggesting that quality of available food resources may influence sex ratio in this species. Mean specific growth rates (mm mm⁻¹ d⁻¹) ranged from 0.013 to 0.016, and matched previous estimates of growth from field populations. Overall ecological performance (survival + growth + reproduction) was similar for all food treatments, except senescent sheaths, which yielded a final mean (±SD) dry biomass (0.4 ± 0.42 mg replicate⁻¹) of amphipods significantly lower than that of other diets (1.7 ± 0.81 to 2.6 ± 0.69 mg replicate⁻¹). Natural diets derived from decomposing cordgrass leaves can fulfill the nutritional requirements of U. spartinophila populations, but variation in initial amounts of living fungal biomass among the five experimental diets only partially explained the responses of amphipods in our experiment. Structural characteristics and variation in rates of fungal occupation within different portions of cordgrass leaves may affect the amphipod's ability to access plant production made available by decomposers. Received: 12 December 1996 / Accepted: 18 December 1996     

Juvenile growth of the polychaete Nereis virens feeding on a range of marine vascular and macroalgal plant sources

A laboratory experiment was conducted to examine variation of juvenile growth (% d^-1) of the polychaete Nereis virens (Sars) in relation to tidal flat plant species as food sources. We used vegetable materials (algae and vascular plants) which are carried along by tidal currents and are found abundantly at the upper tidal level. Juveniles (2-yr-old) markedly increased in wet weight with the algae Laminaria longicruris (weight-specific growth rate : 1.7% d^-1) and Enteromorpha intestinalis (: 1.6% d^-1) as food sources. A higher value of assimilation efficiency was observed for algal species (L. longicruris 55.1±7.9%; E. intestinalis 54.8±0.5%; Fucus vesiculosus 40.6%) than for marine vascular plants (Spartina alterniflora 26.8±10.9%; Zostera marina 1.4%). The digestion of marine vascular species lasts longer (19 to 38 h) than that of algal species. All these characteristics (growth, assimilation efficiency and duration of the digestive process) seem to correlate positively with lignin and cellulose concentrations in the plant structure.     

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.     

Effects of salinity on food intake,absorption and conversion in the rainbow trout Salmo gairdneri

In rainbow trout (Salmo gairdneri Richardson) which had been acclimatised to the experimental salinity and temperature, food intake was maximal in the intermediate salinities of 15.0 and 28.0%, less in fresh water and 7.5% and minimal, by a statistically significant margin, in 32.5%. There were marked day-to-day fluctuations in food intake. When salinity was abruptly increased by 7.5 or 13.0%, there were decreases in growth rate which were related to decreases in food intake. Recovery of food intake and growth rate to pre-increase levels was complete within 14 days. Absorption efficiency, in terms of total dry matter, total energy and total nitrogen, was negatively related to salinity. Total nitrogen was absorbed considerably more efficiently than either energy or dry matter. Conversion efficiency (K ₁ and K ₂) was estimated, also in terms of dry matter, energy and total nitrogen, in trout of the O+ and 1+ year groups weighing from 50 to 150 g. There was a tendency for conversion efficiency to decline with salinity, especially between 28.0 and 32.5%. Dry matter and energy conversion were significantly lower than nitrogen conversion efficiency.     

Effect of delayed first feeding on larval performance of the spider crab Maja brachydactyla assessed by digestive enzyme activities and biometric parameters

The effect of food deprivation on larval performance of the spider crab Maja brachydactyla was studied in terms of survival, moulting capacity, size, weight and enzymatic activities. Five feeding treatments that differed in the initial age of first feeding larvae (fed from hatching, 2, 4 and 6 days post-hatching and unfed) were tested for 20 days. Newly hatched larvae kept without food supply lasted for 10 days and did not moult; with 50% survival observed at 6 days post-hatching. Larvae (zoea I stage) were only able to tolerate 2 days of food deprivation after the onset of exogenous feeding without their performance being compromised. Multivariate analyses suggest that digestive enzyme activities may be good indicators of the nutritional condition of larvae.     

Relationships between porewater nutrients and seagrasses in a subtropical carbonate environment

The feeding, diet and egg production of the copepod Acartia tonsa were dermined during ten experiments in Los Angeles Harbor, California, between November 1986 and October 1987. Copepods were incubated in situ, in quasi-natural food environments. Water temperatures ranged from 14.6 to 21.5°C. Particulate organic carbon and nitrogen (POC and PON) were high (534 to 3710 g Cl^-1, 51 to 459 Nl^-1) but dominated by small (<8 m diam) particles. Plankton (phytoplankton and microzooplankton) C-biomass composed about 10% of the total POC and was usually dominated by particles >8 m. Plankton biomass was always low. Daily ingestion rates ranged from 3 to 96% of body C; egg production ranged from 4 to 35% of body carbon. Mean ingestion and egg production rates during spring-summer were 1.9 and 1.5 times higher than average for the entire study, respectively. The average gross efficiency of egg production for the study was 80%; the spring-summer mean was 41%. Bivariate and multiple-regression analyses revealed that the ingestion rate was dependent upon both temperature and food availability, but that, below 21°C, egg production depended more upon temperature than upon food concentration. To detect dietary preferences, the composition of diet was compared with that of the food supply. Selective feeding was infrequent, but the diet was often dominated by dinoflagellates and ciliates. It would appear that within metabolic limits governed by temperature, the feeding response of A. tonsa is dependent upon food concentration, while egg production depends more on qualitative attributes of the food supply.     

“Good” and “bad” diatoms: development,growth and juvenile mortality of the copepod <Emphasis Type="Italic">Temora longicornis</Emphasis> on diatom diets

We measured development, growth and juvenile mortality of the common copepod Temora longicornis on 11 different monospecific diatom diets in order to estimate (1) how common the negative effects of diatoms are on the development of this copepod and (2) whether the arrested development is connected to deleterious polyunsaturated aldehydes (PUA) or food nutritional quality. Four diatom species (Thalassiosira weissflogii, Thalassiosira rotula CCMP1647, Leptocylindricus danicus CCPM469 and Skeletonema costatum CCMP1281) supported complete development, whereas development failed in or before metamorphosis on seven diatom species/strains (Chaetoceros affinis CCMP158, C. decipiens CCMP173, C. socialis, T. rotula CCMP1018, Thalassiosira pseudonana CCMP1010 and CCMP1335). However, four out of these seven species were not ingested by nauplii, either due to morphology (Chaetoceros spp.) or large size (T. pseudonana CCMP1010). The growth rate did not correlate with the ingestion rate of PUA, neither with ingestion of food mineral (nitrogen) nor with biochemical (polyunsaturated fatty acids, sterols) components. We show that, although some diatoms are of inferior food quality, this is unlikely to be connected to toxicity or due to a direct limitation by a single food nutritional compound.     

What describes fecundity of Capitella sp. 1 better: macro- or micronutrient availability?

During a preliminary set of experiments (April 1987). I measured short-term (20 d) reproductive responses of laboratory-cultured Capitella sp. 1 at 20°C, at four daily ration levels (0.26 to 2.60 mg of organic nitrogen per bowl), and for four food types: Gerber's mixed cereals, Ulva sp., diatoms, and Tetramen fish food. Average female body size, number of fertile segments, and fecundity were significantly affected both by food type and food ration. Average fecundities ranged from 21 (Gerber, 0.26 mg N d^-1) to 448.3 eggs per female (Tetramen, 2.60 mg N d^-1). During a second set of experiments (May 1987), I measured the reproductive response of laboratory-cultured Capitella sp. 1 at 20°C at six ration levels of spring and summer sediment-trap material collected in the Patuxent Estuary, Maryland, USA. Worms raised on these two diets failed to reproduce during the period of this experiment. I used simple and miltiple linear-regression models to describe relationships between fecundity and daily rations and macro- (dry wt, C, N) and micronutrients (essential amino- and essential fatty acids). These models were carried out on three data sets corresponding to either 4 or 6 food types. The results suggest that different nutritional factors may be limiting for different foods. However micronutrients and especially essential amino acids probably more aptly describe nutritional limitation of reproduction in Capitella sp. 1.     

Activity of the principal digestive enzymes in the detritivorous apodous holothuroid Leptosynapta galliennei and two other shallow-water holothuroids

Digestive ability by means of hydrolytic enzyme activities was studied in Leptosynapta galliennei (Herapath, 1865) and two other shallow-water deposit-feeding holothuroids: L. inhaerens (O. F. Müller, 1776) and Holothuria (Panningothuria) forskali (Delle Chiaje, 1828). The holothuroids were collected throughout the year at Roscoff (Britanny, France) from 1980 to 1985, except for L. inhaerens, which was collected in Galway Bay (Ireland) in June/July 1980. Using 25 substrates, the activity of 21 enzymes were recorded to detect hydrolysis of esters (5 substrates), glycosidic bonds (12 substrates) and peptide bonds (8 substrates). The activities of the homogenates of various parts of the gut as well as the digestive juice were measured. Esters and relatively long-chained fatty acids (up to C₁₄) were hydrolysed. Significant hydrolysis of naturally occurring disaccharides and starch (reserve carbohydrate) occurred. The holothuroids were also able to digest hydrolysis products of cellulose and chitin. This may also indicate a capacity to digest the products of complex compounds such as glycolipids or glyco-proteins. Exopeptidases capable of hydrolyzing peptides were present, but endopeptidases able to digest proteolytic chains were not. Most enzymatic activities occurred in all parts of the gut, but some were strictly localized, usually to the anterior intestine. Extracellular digestion of fatty acids, starch and saccharose occurred only in this latter digestive segment. Disaccharidases and peptidases are linked to the plasma membrane of enterocytes. Enzymatic activities associated with lysosomes occur throughout the gut. The diet of shallow-water holothuroids is omnivorous, with a marked preference for food of vegetable origin. Annelids and holothuroids possess a similar capacity to hydrolyse most types of bonds, including peptide bonds. The chemical nature of detritus and its relationship to the nutritional requirements of detritivores must be defined, and gut-retention time ascertained, to discover if different digestive strategies are developed by different detritivores to exploit the same food source in a given environment.     

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