The fish community of the Swartvlei estuary and the influence of food availability on resource utilization

Structural and functional characteristics of the Swartvlei estuary fish community are described. The detritivore group of fishes comprised 49% of the total catch biomass, zoobenthivores 25%, herbivores 18%, piscivores 6%, and epifauna/zooplanktivores 2%. The diets of 18 fish species, based on the analysis of 1,648 stomach contents, are presented. Selection by Swartvlei estuary fishes for zoobenthic invertebrates and aquatic macrophytes in different habitats was calculated using the linear index of selection. Results indicated a strong positive selection for epifaunal invertebrates and poor utilization of infauna and plants. Plant consumption by herbivorous fishes in the estuary centered around filamentous algae and diatoms growing onZostera capensis, rather than seagrass leaf material. The detrital base for both eelgrass- and sand-dominated areas in the Swartvlei estuary is emphasized, and most of the fish biomass was supported directly or indirectly by detritus. Large catches (mass per unit effort) in the macrophyte-free upper reaches site were attributed to the tidal input ofZostera leaves and associated algae to the area and its subsequent utilization by fishes through the detrital food chain.     

Epiphyte-grazer relationships in seagrass meadows: Consequences for seagrass growth and production

Studies of seagrass meadows have shown that the production of algal epiphytes attached to seagrass blades approaches 20% of the seagrass production and that epiphytes are more important as food for associated fauna than are the more refractory seagrass blades. Since epiphytes may compete with seagrasses for light and water column nutrients, excessive epiphytic fouling could have serious consequences for seagrass growth. We summarize much of the literature on epiphytegrazer relationships in seagrass meadows within the context of seagrass growth and production. We also provide insights from mathematical modeling simulations of these relationships for a Chesapeake BayZostera marina meadow. Finally we focus on future research needs for more completely understanding the influences that epiphyte grazers have on seagrass production.     

Conservative mixing of stable isotopes across estuarine salinity gradients: A conceptual framework for monitoring watershed influences on downstream fisheries production

Strong changes in stable isotope tracers commonly occur across estuarine salinity gradients from freshwater to the sea. The tracer gradients reflect the different geochemistries and mixing of freshwater and seawater, and these bottom-up geochemical influences are recorded in estuarine food webs in the isotopic compositions of animals. Conservative mixing calculations suggest that watershed-level inputs of freshwater and nutrients should exert strong influences on isotopic values of estuarine consumers, especially consumers such as bivalves that largely depend on phytoplankton production. Deviations from conservative isotope mixing also occur, and the magnitude of these deviations measures the strength of within-estuary organic matter cycling for estuarine food webs, especially inputs of non-phytoplankton foods such as macrophyte detritus and benthic algae. Measuring consumer isotopes across salinity gradients should be a relatively simple way to monitor effects of watershed nutrient loading and hydrologic flushing in supporting estuarine fisheries production.     

Food Web Structure in a Chesapeake Bay Eelgrass Bed as Determined through Gut Contents and <Superscript>13</Superscript>C and <Superscript>15</Superscript>N Isotope Analysis

Changes in seagrass food-web structure can shift the competitive balance between seagrass and algae, and may alter the flow of energy from lower trophic levels to commercially important fish and crustaceans. Yet, trophic relationships in many seagrass systems remain poorly resolved. We estimated the food web linkages among small predators, invertebrate mesograzers, and primary producers in a Chesapeake Bay eelgrass (Zostera marina) bed by analyzing gut contents and stable C and N isotope ratios. Though trophic levels were relatively distinct, predators varied in the proportion of mesograzers consumed relative to alternative prey, and some mesograzers consumed macrophytes or exhibited intra-guild predation in addition to feeding on periphyton and detritus. These findings corroborate conclusions from lab and mesocosm studies that the ecological impacts of mesograzers vary widely among species, and they emphasize the need for taxonomic resolution and ecological information within seagrass epifaunal communities.     

Role of larger herbvores in seagrass communities

The nutritional ecology of macroherbivores in seagrass meadows and the roles of grazing by urchins, fishes and green turtles in tropical systems and waterfowl in temperate systems are discussed in this review. Only a few species of animals graze on living seagrasses, and apparently only a small portion of the energy and nutrients in seagrasses is usually channeled through these herbivores. The general paucity of direct seagrass grazers may be a function of several factors in the composition of seagrasses, including availability of nitrogen compounds, presence of relatively high amounts of structural cell walls, and presence of toxic or inhibitory substances. The macroherbivores, however, can have a profound effect on the seagrass plants, on other grazers and fauna associated with the meadow, and on chemical and decompositional processes occurring within the meadow. Grazing can alter the nutrient content and digestibility of the plant, as well as its productivity. Removal of leaf material can influence interrelations among permanent and transient faunal residents. Grazing also interrupts the detritus cycle. Possible consequences of this disruption, either through acceleration or through decreased source input, and the enhancement of intersystem coupling by increased export and offsite fecal production, are discussed. The extent and magnitude of these effects and their ecological significance in the overall functioning of seagrass meadows only can be speculated, and probably are not uniform or of similar importance in both tropical and temperate seagrass systems. However, areas grazed by large herbivores provide natural experiments in which to test hypotheses on many functional relations in seagrass meadows.     

海草床食物链有机碳传递过程的研究进展

海草床是重要的近海生态系统,生产力极高,可为海洋动物提供良好的栖息地和丰富的有机碳食源;人类活动引起近岸海域的富营养化,可能会改变海草床有机碳源组成和性质,进而影响植食动物和次级消费者的摄食过程及其食物链能量传递效率,从而影响海草床的生物资源产出功能。归纳总结了国内外海草床食物链碳传递过程各方面的研究进展,主要包括:①海草床有机碳源组成及食源贡献;②海草床初级消费者对有机碳源的摄食;③海草床食物网营养级结构及其能量传递;④富营养化对海草床有机碳源组分结构及其成分的影响;⑤海草床食物链能量传递效率对富营养化的响应。在此基础上,提出了未来的研究重点:①联合运用总有机物的稳定同位素比值法、脂肪酸标志法和特定化合物稳定同位素分析方法,加强定量研究海草床各有机碳源的贡献、食物网营养结构图谱、关键碳流途径及其季节性变化规律;探究关键消费者成体和幼体的有机碳源差异,弄清生长阶段的食性转化过程。②结合野外观测、原位围隔实验和室内模拟实验,深入探讨富营养化对海草床有机碳源组成(海草与附生藻类等)及其化学成分(营养质量和次生化合物)的影响,海草等初级生产者成分变化对植食动物和次级消费者摄食过程的影响,以及牧食食物链和碎屑食物链的组成和碳传递效率等的响应机制。     

Research Progress of the Transfer Process of Organic Carbon Through Food Chain in Seagrass Bed

Seagrass beds are highly productive coastal ecosystems, which provide good nursery habitat and abundant Organic Carbon Sources (OCS) as food for marine animals. Human activities have led to widespread eutrophication in coastal areas. Eutrophication may alter the composition and properties of OCS, thereby affecting the feeding process of herbivores and secondary consumer, and energy transfer efficiency in food chain. This may affect the production function of biological resource in seagrass beds. Based on the summary of the foreign and domestic researches, primary achievements were systematically reviewed in this paper in five aspects: the composition of OCS and their contribution, feeding process of herbivore, food web structure and energy transfer efficiency, and their responses to eutrophication. Future researches that should be emphasized were also prospected. With the combined application of stable isotope analysis for bulks and tissues, fatty acid biomarkers and compound-specific stable isotope analysis, the quantitative study of the contribution of OCS, food web structure, key carbon flow pathway and their seasonal change patterns should be enhanced. Meanwhile, the differences of OCS between larval and adult stages of key consumers will need to be further examined to clarify the transformation of their feeding habits. Through field investigation, in-situ mesocosm and laboratory simulation experiments, the effects of nutrient increase on the structure (seagrass and epiphyte, etc.) and chemical composition (nutritional quality and secondary compounds, etc.) of OCS, the response of feeding process of herbivore and secondary consumer to the alteration of chemical compounds in primary producers, and the response mechanism of carbon transfer efficiency of the grazing food chain and detritus food chain need to be further studied.     

The effect of nitrogen loading on a brackish estuarine faunal community: A stable isotope approach

Coastal ecosystems worldwide face increased nutrient enrichment from shoreline and watershed development and atmospheric pollution. We investigated the response of the faunal community of a small microtidal estuary dominated byRuppia maritima (widgeon grass) in Maine, United States, to increased nitrogen loading using an in situ mesocosm enrichment experiment. Community response was characterized by assessing quantitative shifts in macroinvertebrate community composition and identifying changes in food web structure using stable carbon and nitrogen isotope ratios of producers and consumers. The community was dominated by brackish water invertebrates including midge larvae, oligochaetes, damselfly larvae, amphipods, and ostracods. Experimental nutrient additions resulted in significantly lower densities of herbivorous chironomids and predatory damselflies and greater densities of deposit feeding oligochaetes. Grazing midge larvae (Chironomidae:Dicrotendipes, Cricotopus) consumed epiphytic algae under both natural and enriched conditions. Deposit feedingChironomus was dependent on allochthonous sources of detritus under natural conditions and exhibited a shift to autochthonous sources of detritus under enriched conditions. PredatoryEnallagma primarily consumed grazing chironomids under all but the highest loading conditions. Experimental nutrient loading resulted in an increase in generalist deposit feeders dependent on autochthonous sources of detritus.     

Detritus fuels ecosystem metabolism but not metazoan food webs in San Francisco estuary's freshwater delta

Detritus from terrestrial ecosystems is the major source of organic matter in many streams, rivers, and estuaries, yet the role of detritus in supporting pelagic food webs is debated. We examined the importance of detritus to secondary productivity in the Sacramento and San Joaquin River Delta (California, United States), a large complex of tidal freshwater habitats. The Delta ecosystem has low primary productivity but large detrital inputs, so we hypothesized that de tritus is the primary energy source fueling production in pelagic food webs. We assessed the sources, quantity, composition, and bioavailability of organic matter among a diversity of habitats (e.g., marsh sloughs, floodplains, tidal lakes, and deep river channels) over two years to test this hypothesis. Our results support the emerging principle that detritus dominates riverine and estuarine organic matter supply and supports the majority of ecosystem metabolism. Yet in contrast to prevailing ideas, we found that detritus was weakly coupled to the Delta's pelagic food web. Results from independent approaches showed that phytoplankton production was the dominant source of organic matter for the Delta's pelagic food web, even though primary production accounts for a small fraction of the Delta's organic matter supply. If these results are general, they suggest that the value of organic matter to higher trophic levels, including species targeted by programs of ecosystem restoration, is a function of phytoplankton production.     

The seagrass skeletal assemblage from modern to fossil and from tropical to temperate: Insight from Maldivian and Mediterranean examples

Seagrasses are marine angiosperms that form extensive submarine meadows in the photic zone where carbonate producing biota dwell as epiphytes on the leaves or as infaunal forms, and act as prolific carbonate sediment factories. Because seagrasses have a low preservation potential and records of exceptionally well‐preserved and plant material from marine settings are rare, these palaeoenvironments are difficult to identify in the rock record. Consequently, sedimentological and palaeontological proxies are the main indicators of the presence of seagrass‐dominated ecosystems. This work investigates the skeletal assemblage of Modern (Maldivian and western Mediterranean) and fossil (Eocene; Apula and Oman carbonate platforms and Oligocene; Malta platform) seagrass examples to characterize the skeletal assemblage of modern and fossil seagrasses. Two main types of grains, calcareous algae and foraminifera, constitute around 50% of the bioclastic sediment in both tropical Maldivian and temperate Mediterranean scenarios. However, in the tropical setting they are represented by green algae (Halimeda), while in the Mediterranean they are represented by corallinacean red algae. In contrast, in the Eocene examples, the foraminifera are the most conspicuous group and the green algae are also abundant. The opposite occurs in the Maltese Chattian, which is dominated by coralline algae (mean 42%), although the foraminifera are still abundant. It is suggested to use the term foralgal to identify the seagrass skeletal assemblage. To discriminate between red algae and green algae dominance, the introduction of the prefixes ‘GA’ (green algae) and ‘RA’ (red algae) is proposed. The investigated examples provide evidence that the green algae–foralgal assemblage is typical of tropical, not excessively dense seagrass meadows, characterized by a well‐illuminated substrate to support the development and calcification of the Halimeda thallus. Contrarily, the red algae‐foralgal assemblage is typical of high density tropical to subtropical seagrass meadows which create very dense oligophotic conditions on the sea floor or in temperate settings where Halimeda cannot calcify.     

Variations in stable carbon isotope ratio of the copepod Acartia tonsa during the onset of the Texas brown tide

The Laguna Madre of South Texas is a shallow coastal lagoon whose dominant primary producers shifted from seagrasses to phytoplankton with the onset of the Texas brown tide, which persisted from 1990 through 1997. Acartia tonsa is the dominant component of the mesozooplankton and forms an important link in both the phytoplankton and detritus-based pelagic food webs. Stable carbon isotope ratios of A. tonsa, as well as the two major primary producers: phytoplankton (as particulate organic carbon) and seagrasses, were measured from March 1989 to October 1991. Zooplankton samples were collected at four locations in the Laguna Madre: two in shallow water (c. 1 m) over seagrass beds and two in slightly deeper water (c. 2–3 m) over a muddy bottom in a secondary bay without seagrasses. We found seasonal trends in the isotopic composition of A. tonsa collected within both habitats as well as distinct differences between the average {ie995-1} values of individuals collected in the two regions. Isotopic ratios of animals collected during the summer months were generally 4–8‰ enriched in ¹³C compared with those collected in the winter, at all stations. A. tonsa collected over seagrass beds were 2–5‰ more enriched in ¹³C than those collected over muddy bottoms. These observations suggest carbon derived from seagrasses can be an important source of nutrition for these copepods in summer, especially for copepods living over seagrass beds. The effects of the persistent brown tide decreased the contribution of seagrasses as a carbon source for A. tonsa during the summer of 1991. The pathway by which seagrass carbon enters the diet of A. tonsa is unclear, but the two pathways considered most likely are through copepods feeding on microzooplankton that have fed on bacteria nourished on seagrass carbon, or by copepods feeding directly on particles of seagrass detritus.     

Lessons learned: The effects of nutrient enrichment on the support of nekton by seagrass and salt marsh ecosystems

Coastal ecosystems such as eelgrass beds and salt marshes have always been valued for their high productivity and rich bounty of fish and shellfish. High plant productivity, complex physical structure, and suitable environmental characteristics combine to create areas of high production of important recreational and commercial species. If we are to successfully manage and restore these ecosystems, it is important to understand the mechanisms by which support of nekton may be affected by nutrient enrichment. A review of the literature suggests that there are some similarities and differences in the effects of nutrient enrichment on the support of nekton by seagrass and salt marsh ecosystems. Nutrient enrichment may compromise the ability of these habitats to support fish and invertebrates before the habitat itself is gone. In both ecosystems, alteration of characteristics within the ecosystem (for example, stem density in seagrass and food webs in marshes) affect the support of nekton, even though the basic ecosystem is still clearly extant. Because of differences in natural ecosystem characteristics, loss of ecosystem function does not occur through the same mechanisms. In seagrass systems, physical structure is usually lost first, followed by alteration of food webs and finally changes in dissolved oxygen. In salt marsh systems, loss of dissolved oxygen may occur early in the process, followed by food web alterations and eventually changes in the physical structure may occur. For both seagrass and salt marsh ecosystems, the mechanisms suggested to operate at the ecosystem-level are often based on relatively small-scale plot experiments that have been conducted in only a few locations. A better understanding of how these ecosystems function across broad geographic regions will be needed to ensure functioning coastal ecosystems.     

Dependence of fishery species on salt marshes: The role of food and refuge

Salt marshes are widely believed to serve as nurseries for many fishes and crustaceans of fishery value as a result of the high production of vascular plant detritus and the protection from predation offered by shallow, spatially complex habitats. Comparisons of the yields of species which reside in salt marsh habitats during critical life history stages (such as penaeid shrimp) with the area of such habitats and their greater densities in flooded marshes and associated tidal creeks support the premise that marshes enhance the yield of such species. A range of evidence, including the amount of detrital export from marshes, the poor nutritive value of vascular plant detritus, and natural diets, casts doubt on the notion that production of fishery species is based on the direct consumption of marsh grass detritus or predominantly on food chains based on this detritus. Vascular plants and associated algae may, however, contribute to the production of prey species. The limited observations available support the hypothesis that salt marshes offer significant escape from mortality due to predation, but there have been yet few experimental tests of this hypothesis. Knowledge of relative importance of the food and refuge functions in support of living resources is of practical value in marsh and fisheries management. Better understanding of these roles is important to the effective evaluation of the effects of coastal habitat modifications on fisheries resources and design of alterations to minimize the losses of these values.     

Dynamics of epiphytic photoautotrophs and heterotrophs inZostera marina (eelgrass) microcosms: Responses to nutrient enrichment and grazing

The combined effects of nutrient enrichment and grazing by isopods and amphipods on abundances of seagrass epiphytes were tested inZostera marina L. (eelgrass) microcosms. Using epifluorescence microscopy, densities of epiphytic diatoms, cyanobacteria, heterotrophic flagellates, and heterotrophic bacteria were enumerated after 1 mo and 2 mo of treatment. In general, numbers of diatoms decreased, in the presence of grazers and showed little response to nutrient enrichment, whereas numbers of cyanobacteria increased with nutrient enrichment and showed little response to grazing. Thus, macrofaunal grazing maintained a photoautotrophic community domainated by cyanobacteria, particularly under nutrient enriched conditions. Following 2 mo of treatment, dense macroalgal growth under nutrient-enriched conditins with grazers absent appeared to limit populations of both epiphytic autotrophs. Patterns of abundance of heterotrophic bacteria suggested that the original bacteria population was nutrient limited. Bacteria populations may have been limited by organic carbon supplies at the end of the experiment. Abundances of heterotrophic flagellates and bacteria were strongly correlated on both sampling dates. Results suggest that heterotrophic flagellates might serve as a link between heterotrophic bacterial production and higher trophic levels in seagrass epiphyte food webs.     

Seasonal variation in food web composition and structure in a temperate tidal estuary

Seasonal variation in aquatic food web structure at Mad Island Marsh, Matagorda Bay, Texas, was examined using dietary information obtained from the analysis of gut contents from large samples of fish and crustacean specimens. Unique aspects of this study include the use of large samples of consumer gut contents (n=6,452), long-term sampling (bimonthly surveys over 18 mo), and standard methods of data collection and analysis facilitating comparisons with other aquatic food webs. Dietary data were partitioned for analysis into warm (summer) and cold (winter) seasons. Most consumers fed low in the food web, with trophic levels ranging from about 2 to 3.5 during both summer and winter. Vegetative detritus was more important in macroconsumer diets than live algae and macrophytes. Low trophic levels of consumers reflected the important role of abundant detritivores (e.g., striped mulletMugil cephalus, Gulf menhadenBrevortia patronum, and macroinvertebrates) in linking detritus to top predators via short food chains, a finding consistent with many other estuarine food web studies. Despite changes in community composition and population size structure of certain species, most food web properties revealed comparatively little seasonal variation. The summer food web had more nodes (86), more links (562), a higher density of links as indicated by connectance (0.08), and a slightly higher predator: prey ratio (0.51) compared to the winter food web (75 nodes, 394 links, connectance = 0.07, predator: prey ratio = 0.47). Proportions of top (0.06–0.07), intermediate (0.75–0.76), and basal (0.19) species did not vary significantly between seasons, but mean trophic level was higher during summer. Addition of feeding links based on information from the literature increased connectance to 0.13 during both seasons; other web parameters had values similar to those obtained for our directly estimated food webs. Seasonal variation in food web structure was influenced by changes in community composition (e.g., influxes of postlarval estuarine-dependent marine fishes during winter), availability of resources (e.g., more submerged macrophytes and amphipods during summer), and size structure and ontogenetic diet shifts of dominant consumer taxa. Our findings suggest that some basic properties of estuarine food web are resilient to seasonal changes in population and community structures and food web architecture.     

Faunal communities in seagrass beds: A review of the influence of plant structure and prey characteristics on predator-prey relationships

When compared with nearby unvergetated areas, seagrass meadows contain a dense and strikingly rich assemblage of vertebrates and invertebrates. Most recent literature has focused on evaluating the role of predation in structuring seagrass faunal communities; however, habitat complexity, abundance of food and sediment stability may also be important. This paper summarizes studies relating predator-prey relationships to different features of the seagrass system. This review suggests that the abundance of many species, both epifauna and infauna, is positively correlated with two distinct aspects of plant morphology: 1) the root-rhizome mat, and 2) the plant canopy. A scheme was developed that defines the conditions under which any particular species will be abundant or rare in a seagrass assemblage. This scheme is based on prey and predator characteristics (e.g., epifaunal vs. infaunal, tube-dweller vs. nontube dweller, burrowers vs. nonburrowers, and large vs. small as adult) and on characteristics of the seagrasses (e.g., leaf morphology, shoot density, shoot biomass, structural complexity of the meadow, and root-rhizome density and standing crop).     

Gut contents of common mummichogs,Fundulus heteroclitus L., in a restored impounded marsh and in natural reference marshes

We examined the gut contents of mummichogs, Fundulus heteroclitus L., entering and leaving ditches in three marsh regions within the Barn Island Wildlife Management Area in Connecticut: a restored impounded valley marsh, a marsh below the impoundment dike (Headquarters Marsh), and an unimpounded valley marsh (Davis Marsh). On the Headquarters Marsh and at certain times on the other two marshes, fish entered the ditches on the flooding tide with relatively little food in their guts and left them on the following ebbing tide with considerably more food in their guts. Since the high tides did not flood the surface of the high marsh, it appears that the ditches are important foraging areas. Major components of the gut contents were detritus, algae, amphipods, tanaids, copepods, and insects. During the summer, fish in the restored impounded marsh consumed less food per unit body weight than did fish inhabiting the other marsh regions.     

Fish assemblages in natural versus well-established recolonized seagrass meadows

Studies of fish assemblages between natural and newly recolonized (<4 yr) seagrass meadows have shown no significant differences in community composition between meadow types. However, comparison of natural and well-established (31 yr) recolonized seagrass meadows in the Indian River Lagoon, Florida, showed that, although patterns in fish assemblages are complex and not always consistent, differences were evident. Species richness was higher in natural meadows during spring and autumn while density and species richness were higher in recolonized meadows during summer. Juveniles of all but the five most abundant species were more common in one or the other meadow type. Additionally, species composition was distinctly different between recolonized and natural seagrass meadows, as indicated by UPGMA cluster analysis based on the Morisita-Horn similarity index, Spearman'sr _s (r _s>0.05 in all but one case), and a maximum of only 58.5% species in common. There were also significant differences in the length-frequency distribution for six of seven abundant species. Our results suggest that a well-established recolonized seagrass meadow has the potential to maintain species complements distinct from nearby natural meadows. Reasons for our differing results may include differences in seagrass morphology and collecting techniques between our study and the former studies. Additionally, species may have a longer time to establish specific habitat-use patterns in well-established compared to newly-formed recolonized meadows. Recolonized seagrass meadows appear to be as suitable a habitat as natural meadows for juvenile and small adult fishes.     

A comparison of methods for the accurate measurement of epiphytic carbonate

The accurate measurement of epiphytic carbonate is vital in the construction of carbonate budgets. Prior studies have produced estimates of epiphytic carbonate production, but the results of these investigations are not directly comparable due to the variety of methods employed by the investigators. This investigation field-tested four methods of measuring epiphytic carbonate. These four methods were the gravimetric leaf acidification technique, the EDTA titration of epiphytized leaves, the EDTA titration with seagrass blanks, and the scrape and titrate method. The leaf acidification method and the EDTA titrimetric method without seagrass blanks resulted in significantly larger estimates of epiphytic carbonate than those produced using the other two methods. It is believed that the leaf acidification technique and the EDTA titration of epiphytized leaves results in overestimates of epiphytic carbonate due to the leaching of organic matter during acidification. The scrape and titrate method, involving the separation of epiphyte material from the seagrass leaves, is believed to most accurately measure epiphytic carbonate.