A comparison of two tubificid oligochaete species as candidates for sublethal bioassay tests relevant to subtropical and tropical regions

Two tubificid oligochaetes, Branchiura sowerbyi and Tubifex tubifex, were evaluated as potential test organisms for sediment bioassays. We attempt to reproduce the sediment bioassay proposed by Reynoldson et al. (1991) using his strain of Tubifex tubifex and his reference sediment and to compare this technique using Branchiura sowerbyi. This species was chosen because it is more common and dominant in tropical and subtropical environments than Tubifex tubifex. Data on survival and reproduction were obtained for both species, and growth estimates were obtained for B. sowerbyi. The sublethal bioassay with T. tubifex confirmed earlier estimates of a test duration of 4 weeks at 22.5 °C. B. sowerbyi cultures can produce usable estimates in 21 days at 30 °C.     

Tubifex tubifex as a link in food chain transfer of hexachlorobenzene from contaminated sediment to fish

Sediments contaminated with poorly water-soluble organic chemicals pose a risk to aquatic food chains. Sediment-associated chemicals can be accumulated by endobenthic, sediment-ingesting invertebrates. Some tubificid species – or other benthic annelids – serve as food for benthivorous fish, which thereby ingest the sediment-borne chemicals and may accumulate contaminant concentrations far higher than from water exposure only, and transfer them to organisms of higher trophic levels. For measurement of biomagnification, a sediment based food chain was developed and established in the laboratory. The two-step food chain included the sediment-dwelling freshwater oligochaete Tubifex tubifex (Müller) as a representative species of benthic infauna. The three-spined stickleback (Gasterosteus aculeatus, Linné), a small teleost fish which often feeds primarily on benthic invertebrates, served as a model predator. Spiked artificial sediment and reconstituted water as the overlying medium were used. Experiments were performed using ¹⁴C-labelled hexachlorobenzene, a hydrophobic pollutant as a model compound. To examine the influence of benthic prey on the bioaccumulation of the test substance in the predator, fish were exposed to spiked water, spiked sediment, pre-contaminated prey organisms, or combinations of these exposure routes. The results of these experiments indicate that for hexachlorobenzene, the presence of contaminated Tubifex tubifex as a food source in combined exposure leads to significantly higher accumulation in fish than exposure to single pathways.     

Sedimentary context controls the influence of ecosystem engineering by bioturbators on microbial processes in river sediments

By modifying the physical environment, ecosystem engineers can have inordinately large effects on surrounding communities and ecosystem functioning. However, the significance of engineering in ecosystems greatly depends on the physical characteristics of the engineered habitats. Mechanisms underlying such context‐dependent impact of engineers remain poorly understood even though they are crucial to establish general predictions concerning the contribution of engineers to ecosystem structure and function. The present study aimed to decrypt such mechanisms by determining how the environmental context modulates the effects of ecosystem engineers (bioturbators) on microorganisms in river sediments. To test the effects of environmental context on the role of bioturbators in sediments, we used mesocosms and recreated two sedimentary contexts in the laboratory by adding a layer of either fine or coarse sand at the top of a gravel‐sand matrix. For each sediment context, we examined how the sediment reworking activity of a bioturbating tubificid worm (Tubifex tubifex) generated changes in the physical (sediment structure and permeability) and abiotic environments (hydraulic discharge, water chemistry) of microorganisms. Microbial characteristics (abundances, activities) and leaf litter decomposition – a major microbially‐mediated ecological process – were measured to evaluate the impact of bioturbation on biotic compartment. Our results showed that the permeability, the availability of oxygen and the activities of microorganisms were reduced in sediments covered with fine sand, in comparison with sediments covered with coarse sand. Tubifex tubifex significantly increased permeability (by about six‐fold), restored aerobic conditions and ultimately stimulated microbial communities (resulting in a 30% increase in leaf litter breakdown rate) in sediments covered with fine sand. In contrast T. tubifex had low effects in sediments topped by coarse sand, where O₂ was already available for hyporheic microorganisms. Our study supports the idea that context dependency mainly modulates the effects of engineering by controlling the ability of engineers to create changes on abiotic (O₂ in the present study) factors that are limiting for surrounding communities.     

Host–parasite interactions and competition between tubificid species in a benthic community

1. Parasites can be important determinants of host community structure while host community structure can influence the success of parasites, although both are often overlooked. In two laboratory experiments, we examined interactions among Myxobolus cerebralis syn Myxosoma cerebralis Höfer, the myxozoan parasite that causes salmonid whirling disease, and two coexisting tubificid species: Tubifex tubifex (Müller), which is the alternate host of the parasite, and Limnodrilus hoffmeisteri Claparède, which is not susceptible. In the first experiment, we examined T. tubifex infection prevalence when exposed to nine doses of spores. In the second experiment, we examined tubificid and parasite success under three spore doses when tubificids were combined in a response surface experimental design used to detect interactions among species. 2. The outcomes of interactions between tubificid species were complex. The number and biomass of offspring of both tubificid species were density dependent when in monoculture or in combination with the other species. Adult growth of T. tubifex was also density dependent in monoculture, but when L. hoffmeisteri replaced one‐half of the T. tubifex in the high‐density treatment, adult growth of T. tubifex was higher than in monoculture. Adult growth of L. hoffmeisteri was always density independent. Whether T. tubifex was exposed to the parasite or not did not change the outcome of these interactions. However, adult growth of T. tubifex, but not L. hoffmeisteri, was highest when M. cerebralis was present. 3. Infection prevalence of T. tubifex increased with increasing spore dose. Infection prevalence was lowest in the high‐density T. tubifex monoculture and highest in the low‐density T. tubifex monoculture and when T. tubifex was in combination with L. hoffmeisteri. 4. Both intraspecific and interspecific competition influenced tubificid success, but T. tubifex gained some competitive advantage through increased adult growth when in combination with L. hoffmeisteri. Whether T. tubifex was exposed to the parasite or not did not change the outcome of the interactions between the tubificid species. 5. The presence of L. hoffmeisteri did not decrease the prevalence of infection in T. tubifex, suggesting that parasite success was unaltered by the presence of this non‐susceptible species.     

Sublittoral and profundal Oligochaeta fauna of the Lake Constance (Bodensee-Obersee)

Fourteen tubificid species and one lumbriculid make up the oligochaete population of the northern half of Lake Constance, a meso- to mesoeutrophic lake. Tubifex tubifex makes up 72% of the total abundance, the clean water species Stylodrilus heringianus only 0.1%. No correlation exists between total worm abundance and organic carbon content of the sediment, only 4% of the organic carbon being used by the worms. A highly significant correlation between particulate organic matter loading and its organic carbon content from 10 tributaries and total oligochaete abundance is demonstrated. The ecological index I_PA is defined and used to assess the biologically relevant trophic status of the lake bed. This integrates information on the percent representation of three groups of worm species classified on atrophic basis and total abundance of the worms. Despite control of the increase in phosphorus loadings progressive eutrophication of sediments was observed after an 8 year interval.     

A morphological reappraisal of Tubifex blanchardi Vejdovský, 1891 (Clitellata: Tubificidae)

Tubifex blanchardi Vejdovský, 1891 is a freshwater tubificid, often living in sympatry with Tubifex tubifex (Müller 1774). Although considered from its discovery as a species on its own, its biological status is debated. During the early seventies T. blanchardi was reduced to a mere form of T. tubifex, as a particular case of polymorphism in chaetal pattern. Using classical histological techniques, microdissections of portions of the male genital apparatus and phalloidin staining of dissected copulatory organs we investigated 163 mixed individuals of T. blanchardi and T. tubifex belonging to sympatric populations from the Lambro River (Milan, Northern Italy). The internal morphology of T. blanchardi is described for the first time. Our results show that T. tubifex and T. blanchardi differ in several characters concerning both their external and internal morphology, and in the fine organization of their copulatory organs. Several independent character sets support the separation of T. blanchardi from T. tubifex, suggesting that it is an independent species. This study also supports the idea that T. blanchardi and T. bergi (Hrabě, 1935), another species closely related to T. tubifex, are not conspecific. The observed morphological differences between allopatric populations of T. tubifex are discussed.     

Fertile sediment and ammonium enrichment decrease the growth and biomechanical strength of submersed macrophyte Myriophyllum spicatum in an experiment

Decline of submersed macrophytes has occurred in eutrophic lakes worldwide. Little is known about effects of nutrient enrichment on biomechanical properties of submersed macrophytes. In a 30-day experiment, Myriophyllum spicatum was cultured in aquaria containing two types of sediment (mesotrophic clay vs. fertile loam) with contrasting water NH₄ ⁺ concentrations (0 vs. 3.0 mg L^?1 NH₄–N). The plant growth, shoot and root morphology, stem biomechanical properties, and stem total nonstructure carbohydrates content (TNC) were examined. The NH₄ ⁺-enriched water, particularly combined with the fertile sediment, caused adverse effects on M. spicatum as indicated by reductions in the growth, stem biomechanical properties (tensile force, bending force and structural stiffness), and TNC content. These results indicate that increased sediment fertility and water NH₄ ⁺-enrichment made the plant more fragile and vulnerable to hydraulic damage, particularly for the upper stem, implying that M. spicatum was prone to uprooting and fracture by hydraulic force, and the broken fragment from parent shoot of M. spicatum might have low-survival potential due to its low-TNC content. This may be a mechanical aspect for the decline of submersed macrophytes and makes it more difficult to restore submersed vegetation in the eutrophic lakes.     

Influence of ecosystem engineers on ecosystem processes is mediated by lake sediment properties

Invasive species may impact biotic community structure, ecosystem processes, and associated goods and services. Their impact may be especially strong when they also serve as ecosystem engineers (i.e. organisms affecting the physical habitat and resources for other species). Dreissenid mussels fill both these roles, having invaded the Laurentian Great Lakes in the late 1980s, and also serve as ecosystem engineers by altering nutrient fluxes and influencing the microbial food web through direct nutrient release and excretion of feces and pseudo‐feces at the water–sediment interface. We conducted laboratory experiments to investigate how the different functional traits of invasive quagga mussels (filtering activity and direct nutrient release) and native chironomid larvae (tube building and bioirrigation) interact with lake sediment of differing organic matter content to influence biogeochemical fluxes and water quality. Our results showed that sediment reworking and ventilation activities by chironomid larvae increased oxygen penetration in the sediment, affecting primarily pore water chemistry, whereas invasive mussels enhanced nutrient releases in the surface water. However, sediment organic matter modulated the effects of ecosystem engineers on system‐level processes; chironomids had a greater influence on sediment reworking and microbial‐mediated processes in organic‐rich sediments, whereas quagga mussels enhanced nutrient concentrations in the overlying water of organic‐poor sediments. These results have management implications, as the effects of invasive mussels on the biogeochemical functioning in the Great Lakes region and elsewhere can alter system bioenergetics and promote harmful algal blooms.     

Cyst formation of Tubifex tubifex (Müller) — an adaptation to survive food deficiency and drought

Adult individuals of Tubifex tubifex (Müller) (Oligochaeta, Tubificidae) could be induced to form cysts in the laboratory. Hiding in cysts, Tubifex tubifex survived a five-month drought period in the field. Encysted Tubifex tubifex had a lower super-cooling-point than mobile individuals. Food shortage is believed to be the main factor in cyst formation.     

Size-related auto-fragment production and carbohydrate storage in auto-fragment of <Emphasis Type="Italic">Myriophyllum spicatum</Emphasis> L. in response to sediment nutrient and plant density

Size-related asexual reproduction of submersed macrophytes is still poorly understood. Here, we investigate how size-related auto-fragmentation in Myriophyllum spicatum L. responds to sediment nutrients and plant density. An experiment was carried out with sediments containing two different nutrient levels and with two levels of plant density. The results show that sediment nutrients and plant density brought about a strong dependency of auto-fragment production and the amount of total non-structural carbohydrate (TNC) storage in auto-fragments on individual plant size (total plant biomass). However, these two factors acted differently on size dependency. Sediment nutrients positively affected auto-fragment production and the amount of TNC in auto-fragments of M. spicatum. High concentrations of sediment nutrients significantly increased these two traits in absolute value and the value relative to plant size. Although the auto-fragment biomass and the amount of TNC in auto-fragments did not differ between density treatments when plant size was considered, the absolute values of these two traits were much larger in the low plant density treatment than in the high plant density treatment, which suggested an indirect negative effect of plant density on the auto-fragmentation of M. spicatum. In addition, higher percentages of large auto-fragments (>100 mg) were produced by plants that grew in nutrient poor sediment and low plant density environment than plants in nutrient rich sediment and high plant density environment. These results do not solely highlight a size-dependent effect, but also a size-independent effect of auto-fragment production and the amount of TNC in auto-fragments of M. spicatum. Furthermore, such size-independent effects can be explained by the significant biomass partitioning differences and the similar TNC-concentrations in auto-fragments under different environmental conditions.     

Density-dependent processes in cohorts of Tubifex tubifex,with special emphasis on the control of fecundity

Laboratory cohort cultures of the tubificid Tubifex tubifex with different initial densities were carried out at 20° C with the condition of unlimited food. The main results were: 1) Intracocoon mortality was 37% of the laid eggs (observation of 689 eggs); 2) The principal bionomic parameters (generation time, r, R₀) appeared to be density dependent; 3) Recruitment was regulated through the percentage of worms that actually attained the ovigerous stage, specific fecundity, and the duration of the egg laying stage, which appeared to be inversely correlated with density.     

Morphological and Chemical Changes Induced by Herbivory in Three Common Aquatic Macrophytes

The Dry Matter Content (DMC), the total phenolic content, the production of new branches and the plant fragmentation were compared in three macrophyte species (Elodea canadensis, Elodea nuttallii and Myriophyllum spicatum) exposed or not to snail herbivory. Grazing significantly reduced the DMC of M. spicatum and E. canadensis, but had no effect on the DMC of E. nuttallii. The phenolic contents of Elodea species were not modified by snail herbivory, whereas that of M. spicatum significantly increased when exposed to grazers. The number of new branches produced by M. spicatum and E. canadensis plants, and the fragmentation of E. canadensis also increased in response to herbivory. Chemical defences are therefore probably constitutive in Elodea and induced in M. spicatum, and morphological changes can be related to species growth form and synthesis of phenolic compounds. (© 2009 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Composition and distribution of bottom oligochaete fauna of a north Italian eutrophic lake (Lake Ledro)

The profundal macroinvertebrates, particularly the oligochaetes, of Lake Ledro (Trento, Italy), that has recently undergone eutrophication, were studied.A statistical approach of random sampling was used to study the distribution and abundance of the oligochaete species. The optimum sample number was calculated from a preliminary sample series. The oligochaete community was made up of five tubificid species, one naidid and one lumbriculid species that on average represents more than 80% of the macrobenthic community. Population density was correlated with depth and decreasing oxygen concentration. The role of Tubifex tubifex as a eutrophic, tolerant species was confirmed; and in fact it was the only species found (although at low density) in the deepest and anoxic zone. No comparable data are available for the lake prior to eutrophication, but these data will be valuable for future comparison once a remediation program for the lake has been implemented.     

A field test of a sediment bioassay with the oligochaete worm Tubifex tubifex (Müller, 1774)

A sediment toxicity test using reproduction in the tubificid oligochaete Tubifex tubifex was tested in 11 geographic locations in the Laurentian Great Lakes where sediments have previously been documented as being contaminated. Samples were taken from both potentially contaminated and clean sites in each of the 11 areas. Using test endpoints such as survival and growth, results from 72 samples were classified using cluster and ordination techniques. Six response groups were identified and four groups of toxic sites determined. Toxic effects ranged from mortality of adult worms to reduced reproduction. Using correlation and discriminant analysis there was good evidence that high concentrations of metals such as copper and lead were responsible for the observed effects at some sites.     

Disappearance of deep profundal zoobenthos in Lake Ikeda, southern Kyushu, Japan, with relation to recent environmental changes in the lake

A benthological survey in a deep caldera, Lake Ikeda, southern Kyushu, Japan, in 1998 revealed that no zoobenthos were found in the deep profundal, although two tubificid oligochaetes, Tubifex tubifex and Limnodrilus hoffmeisteri, and a chironomid, Procladius sp., were distributed in the upper profundal zone. This is the first record of oligochaete composition in the lake. Lake Ikeda had been typically oligotrophic until the 1940s, and zoobenthic assemblages were recorded throughout the profundal bottom in the 1920s and 1970s. Recent disappearance of the deep profundal zoobenthos could be caused by the stagnation of anoxic waters in the hypolimnion, in connection with eutrophication triggered by nutrient loading, as well as change in the thermal circulation system presumably caused by global warming.     

亚热带河口陆基养虾塘水体溶解性碳浓度及沉积物-水界面碳通量时空动态特征

以福建闽江和九龙江河口陆基养虾塘为研究对象,通过野外原位观测和室内模拟培养实验,开展了河口陆基养虾塘养殖期间水体溶解性有机碳(DOC)和溶解性无机碳(DIC)及养虾塘沉积物-水界面碳交换通量变化特征的研究。结果表明:时间变化上,养虾塘水体溶解性碳浓度及沉积物-水界面碳通量在闽江河口呈现8月中旬10月中旬6月中旬的特征,在九龙江河口表现为随养殖阶段推移而增加的趋势;空间变化上,闽江河口养虾塘水体溶解性碳浓度及沉积物-水界面碳通量显著高于九龙江河口;沉积物释放溶解性碳速率与水体溶解性碳浓度呈现显著正相关关系,沉积物碳释放过程是引起养虾塘水体溶解性碳浓度时空变化的重要因素。表明河口区水产养虾塘碳循环研究时需考虑不同形态碳生物地球化学循环的时空差异性。     

Effects of invasive species on plant communities: an example using submersed aquatic plants at the regional scale

Submersed aquatic plants have a key role in maintaining functioning aquatic ecosystems through their effects on the hydrological regime, sedimentation, nutrient cycling and habitat of associated fauna. Modifications of aquatic plant communities, for example through the introduction of invasive species, can alter these functions. In the Sacramento-San Joaquin River Delta, California, a major invasive submersed plant, Brazilian waterweed Egeria densa, has become widespread and greatly affected the functionality of the submersed aquatic plant community. Rapid assessments of the distribution and abundance of this species are therefore crucial to direct management actions early in the season. Given the E. densa bimodal growth pattern (late spring and fall growth peaks), summer assessments of this species may indicate which and where other submersed species may occur and fall assessments may indicate where this and other species may occur in the following spring, primarily because the Delta’s winter water temperatures are usually insufficient to kill submersed aquatic plant species. We assessed community composition and distribution in the fall of 2007 and summer of 2008 using geostatistical analysis; and measured summer biomass, temperature, pH, salinity, and turbidity. In the fall of 2007, submersed aquatic plants covered a much higher proportion of the waterways (60.7%) than in the summer of 2008 (37.4%), with a significant overlap between the seasonal distribution of native and non-native species. Most patches were monospecific, and multispecies patches had significantly higher dominance by E. densa, co-occurring especially with Ceratophyllum demersum. As species richness of non-natives increased there was a significant decrease in richness of natives, and of native biomass. Sustained E. densa summer biomass negatively affected the likelihood of presence of Myriophyllum spicatum, Potamogeton crispus, and Elodea canadensis but not their biomass within patches. Depth, temperature and salinity were associated with biomass; however, the direction of the effect was species specific. Our results suggest that despite native and invasive non-native submersed plant species sharing available niches in the Delta, E. densa affects aquatic plant community structure and composition by facilitating persistence of some species and reducing the likelihood of establishment of other species. Successful management of this species may therefore facilitate shifts in existing non-native or native plant species.     

Relative influence of bioturbation and predation on organic matter processing in river sediments: a microcosm experiment

1. Our objective was to measure the effects of bioturbation and predation on the physical characteristics and biogeochemical processes in river sediments. 2. We investigated the impacts of tubificid worms tested separately and together with an omnivore (Gammarus pulex), which does feed on tubificids, on sediment distribution, water flux, sediment organic carbon, biofilm biomass and microbial activities, and the concentrations of dissolved oxygen, dissolved organic carbon, PO, NO, NO and NH in slow filtration sand–gravel columns. We hypothesised that gammarids, which exploit the top 2–3 cm of the sediment, would modify the impact of worms at the sediment surface. 3. In experiments both with and without gammarids, bioturbation by the tubificids modified both the distribution of surface particles in the sediment column and water flux. In addition, microbial aerobic (oxygen consumption) and anaerobic (denitrification and fermentative decomposition of organic matter) processes in the sediment were stimulated in the presence of tubificid worms. However, G. pulex did not affect either the density or bioturbation activity of the tubificid worms. 4. Bioturbation by the benthos can be a major process in river habitats, contributing to the retention of organic matter in sediment dynamics. The presence of at least one predator had no effect on bioturbation in sediments. In such systems, physical heterogeneity may be sufficient for tubificids to escape from generalist predators, though more specialised ones might have more effect.     

The biology of Oligochaeta from Dorset chalk streams

Fewer species and more individuals of Tubificidae were present in the soft sediments of a polluted ditch than in the relatively unpolluted Bere Stream. Changes in width frequency of natural populations of Tubifex tubifex and Limno-drilus hoffmeisteri throughout the year were similar at both the sites investigated. In laboratory experiments larger populations of Tubifex tubifex developed in fine sediments than coarse. By means of a width-dry-weight relationship it was possible to estimate the weight of the experimental populations.     

Effect of Campsurus notatus on NH+4, DOC Fluxes,O2 Uptake and Bacterioplankton Production in Experimental Microcosms with Sediment‐Water Interface of an Amazonian Lake Impacted by Bauxite Tailings

The aim of this study was to evaluate the influence of Campusurus notatus Eaton 1868 (Ephemeroptera: Polimitarciydae) and the impact of bauxite tailings on ammonium (NH₄⁺) and dissolved organic carbon (DOC) fluxes, oxygen uptake and bacterioplankton production in the sediment‐water interface of Lake Batata, a shallow Amazonian floodplain lake. Mesocosms were constructed from natural and impacted areas of Lake Batata, to reproduce the sediment‐water interface. The cores were incubated with 0 to 2,388 ind m^–2 of Campsurus notatus nymphs, and the changes in NH₄⁺, DOC, O₂ concentration and bacterioplankton production in the overlying water column were measured. Ammonium efflux (F = 9.8, p < 0.05, multiple regression) and oxygen uptake (F = 11.8, p < 0.05) showed a significant correlation with the density of C. notatus in the cores with natural sediment. No differences on DOC release were observed in cores with natural or impacted sediment. In the cores incubated with natural sediment and nymphs of C. notatus, a significant increase (Two‐way ANOVA, p < 0.05) in bacterial production (0.44 μg C l^–1 h^–1) was observed after 3 hours of incubation. In cores incubated with sediment impacted by bauxite tailings, there was no difference in bacterial production with and without C. notatus. We conclude that C. notatus is an important bioturbator in Lake Batata, increasing the turnover rate of nitrogen (NH₄⁺) at the sediment‐water interface and bacterial production in cores incubated with natural sediment. It is also clear that bauxite tailings reduce the nutrients turnover rates in impacted regions of Lake Batata and influence bacterial production.