Sexual maturity in young Dreissena polymorpha from Lake Como (N. Italy)

Summary

The reproductive biology of the zebra mussel Dreissena polymorpha was studied during 1999 in a natural population living in Lake Como (N. Italy), with particular reference to the differences in behavior between young and adult mussels. Histological analysis of gonads was used to study the gametogenic cycles and to evaluate when the young reached sexual maturity. Signs of active gametogenesis were detected in ≥4 mm shell length mussels. Mature gametes and the first spawning events were observed in ≥5 mm males and in ≥6 mm females. The gonad feature of ≥7 mm mussels was similar to that of the adults. Size frequency distributions revealed that the new generation (0⁺) first spawned in the year following birth, but only after winter. The length of larval time seemed to vary greatly depending on the spawning period, in addition to the environmental conditions. The first settled mussels ≥500 μm in shell length were observed 3–4 months after the first depositions. Water temperature seemed to be the most important factor which regulated sexual phenomena of both young and adult mussels. Its role in the onset of the spawning events was confirmed, but it is important to point out that temperature is not enough to trigger reproduction since gamete release is impossible without sufficient energy, especially for young mussels.     

In situ survival and growth of zebra mussels (Dreissena polymorpha) under chronic hypoxia in a stratified lake

Experiments and field surveys were conducted in Hargus Lake (Ohio, U.S.) to investigate the effect of lake stratification on the survival, growth and distribution of zebra mussels. During the lake stratification period, relatively stable temperature and dissolved oxygen (DO) gradients persisted across the water column, allowing us to examine the chronic effect of hypoxia on zebra mussels. Zebra mussels were incubated in cages and suspended at different depths in the water column at both pelagic (max. depth = 12 m) and littoral (max. depth = 3.5 m) sites from April 18 to September 28, 1994. No mussel survived to the end of the experiment in cages ≥ 5.5 m, whereas the highest survival rate (76%) occurred at 5 m depth where temperature and DO remained fairly stable for at least 3 months. The threshold oxygen level for survival was between 1.0–1.7 mg l^-1 when water temperature was at about 17–18 °C. While zebra mussels′ survival rate was not affected under the sublethal hypoxic conditions, their growth was greatly retarded by poor water quality. The field survey showed that the zebra mussels and macrophytes had about the same distribution and their biomasses were positively related. The percentage of mussels in aggregates increased towards their maximum distribution depth. The maximum distribution depth of the naturally occurring zebra mussels was only 2.8 m, whereas the adult mussels could survive the entire stratification period when being artificially placed on the 3.5 m bottom, and young mussels could colonize the 3.5 m bottom if solid substrates were provided. We conclude that lack of substrate, rather than hypoxia, was the limiting factor of zebra mussel distribution above 5 m depth in Hargus Lake. This revised version was published online in July 2006 with corrections to the Cover Date.     

Habitat engineering by the invasive zebra mussel <Emphasis Type="Italic">Dreissena polymorpha</Emphasis> (Pallas) in a boreal coastal lagoon: impact on biodiversity

Habitat engineering role of the invasive zebra mussel Dreissena polymorpha (Pallas) was studied in the Curonian lagoon, a shallow water body in the SE Baltic. Impacts of live zebra mussel clumps and its shell deposits on benthic biodiversity were differentiated and referred to unmodified (bare) sediments. Zebra mussel bed was distinguished from other habitat types by higher benthic invertebrate biomass, abundance, and species richness. The impact of live mussels on biodiversity was more pronounced than the effect of shell deposits. The structure of macrofaunal community in the habitats with >10³ g/m² of shell deposits devoid of live mussels was similar to that found within the zebra mussel bed. There was a continuous shift in species composition and abundance along the gradient ‘bare sediments—shell deposits—zebra mussel bed’. The engineering impact of zebra mussel on the benthic community became apparent both in individual patches and landscape-level analyses.     

Reproductive ecology of the striped bitterling Acheilognathus cyanostigma (Cyprinidae: Acheilognathinae)

The reproductive ecology of the striped bitterling Acheilognathus cyanostigma was investigated in a small pond in Mie Prefecture, central Honshu, Japan. A. cyanostigma was the only species of bitterling in the study pond, where only a single species of mussel Anodonta woodiana was present. Spawning of A. cyanostigma was recorded between early April and early July, peaking between late April and mid-June in 2003. Ovipositor length during the spawning period ranged from 19.2 to 42.8 mm, and was positively correlated with female body length, but did not show significant seasonal variation. The eggs are elliptical with a volume of approximately 1.5 mm³. Egg size correlates positively with female body size, and both egg shape and volume changed significantly with season. The embryos were located on host mussel gills approximately 30 mm from the exhalant siphon and were found more frequently on the inner rather than the outer mussel gills. The possible ecological significance of these observations is discussed in the context of the adaptations of A. cyanostigma for utilizing mussels for oviposition.     

Variations in the Reproductive Cycle of Dreissena Polymorpha in Europe, Russia, and North America

SYNOPSIS. The reproductive cycle of the zebra mussel {Dreissenapolymorpha) is highly variable throughout its range in Europe,Russia², and North America. The environmental factors influencingthis variation are poorly understood, but successful reproductionis occurring in areas where it was initially believed that adultzebra mussels could not survive (i.e., southern United States).The differences in mussel reproduction occurring from site-to-sitemake it difficult to predict timing of specific events, suchas the start of larval production, that are important in initiatingcontainment or control procedures. For example, the amount oftime required for a fertilized egg to develop into a juvenilemussel can be as short as 8 days, or as long as 240 days. Releaseof gametes by adults can be a highly synchronized event, focusedover a 1–2 week period, or it can be completely non-synchronized,occurring throughout the year. Zebra mussels in some localitiesstart spawning at water temperatures of 12–13°C, butdo not start until water temperatures reaches 22°C at othersites. While some of this variability in reproductive behaviorstems from mussel adaptation to local conditions, part is dueto difficulties in sampling these events. It is difficult todetermine reproductive success of a specific population becauseof the problems in separating locally produced larvae from larvaedrifting in from other areas. Further research is needed notonly on the relationship between reproduction and environmentat the community level, but also on the variability in responseof individual mussels.     

Seasonal reproductive cycle in the mussel,Lamellidens corrianus

Seasonal reproductive cycle of the freshwater mussel, Lamellidens corrianus has been studied. These mussels are functional or simultaneous hermaphrodites. The spawning was at its peak during the months of September to December. The gonads were in growing stages with reduced gonadal activity during January to April, whereas the maturation of gonads was found to be intense during May to August.     

Effects of increasing temperatures on population dynamics of the zebra mussel <Emphasis Type="Italic">Dreissena polymorpha</Emphasis>: implications from an individual-based model

Zebra mussels (Dreissena polymorpha, Pallas, 1771) have had unprecedented success in colonizing European and North American waters under strongly differing temperature regimes. Thus, the mussel is an excellent model of a species which is able to cope with increasing water temperatures expected under global change. We study three principle scenarios for successful survival of the mussel under rising temperatures: (1) no adaptation to future thermal conditions is needed, existing performance is great enough; (2) a shift (adaptation) towards higher temperatures is required; or (3) a broadening of the range of tolerated temperatures (adaptation) is needed. We developed a stochastic individual-based model which describes the demographic growth of D. polymorpha to determine which of the alternative scenarios might enable future survival. It is a day-degree model which is determined by ambient water temperature. Daily temperatures are generated based on long-term data of the River Rhine. Predictions under temperature conditions as recently observed for this river that are made for the phenology of reproduction, the age distribution and the shell length distribution conform with field observations. Our simulations show that temporal patterns in the life cycle of the mussel will be altered under rising temperatures. In all scenarios spawning started earlier in the year and the total reproductive output of a population was dominated by the events later in the spawning period. For maximum temperatures between 20 and 26°C no thermal adaptation of the mussel is required. No extinctions and stable age distributions over generations were observed in scenario 2 for all maximum temperatures studied. In contrast, no population with a fixed range of tolerated temperatures survived in scenario 3 with high maximum temperatures (28, 30, 32°C). Age distributions showed an excess of 0+ individuals which resulted in an extinction of the population for several thermal ranges investigated. Priority programme of the German Research Foundation—contribution 14.     

Physiological Aspects of Zebra Mussel Reproduction: Maturation, Spawning, and Fertilization

SYNOPSIS. The prolific reproductive capabilities of the zebramussel, Dreissena polymorpha, have facilitated the rapid spreadand high densities of this biofouling organism since its accidentalintroduction into North America less than 10 years ago. Researchon its reproductive mechanisms and capabilities may be valuablenot only in predicting its further spread, but also in investigatingbasic mechanisms of reproduction and development and in developingnew strategies to mitigate its impact. Since zebra mussels aredioecious and fertilization occurs externally, coordinated maturation,spawning, and other mechanisms have evolved to increase theprobability of successful fertilization. The zebra mussel undergoesan annual cycle of gonadal growth and gamete maturation, culminatingin one or more spawning events in late spring or early summer.Temperature, rates of temperature change, food availability,and effects of neighboring mussels seem to be critical variablesthat determine reproductive responses. Serotonin is a biogenicamine which is implicated in spawning behavior and can reliablytrigger spawning. Serotonin is present in the gonad in neuralvaricosities that encircle groups of gametes, and specific serotonergicligands can mimic or block spawning caused by serotonin. Infemales, serotonin reinitiates meiosis causing maturation fromprophase I to metaphase I prior to spawning. Spawned oocytescontain substances that are species specific sperm chemoattractants.The sequence of binding, entry, and subsequent nuclear movementshave been observed with fluorescence and scanning microscopy.Despite their negative ecological and economic impacts, zebramussels have also provided a new and easily obtainable resourcefor studies of reproductive mechanisms.     

Seston quality controls zebra mussel (Dreissena polymorpha ) energetics in turbid rivers

Feeding processes and energetic balance of zebra mussels were both related to the quantity and quality of natural seston. Filtration rate and pseudofeces production increased while clearance rate remained constant with increasing seston concentration. Ingestion rate, assimilation efficiency, and assimilation rate all increased with increasing food quality, measured as the ratio of organic to inorganic material in the seston. Respiration rate did not change with either food quantity or quality. As a result, scope for growth declined with decreasing food quality, and fell below 0 cal mg⁻¹ h⁻¹ at an organic:inorganic ratio of 0.5. The association between feeding processes and food quality appears related to a breakdown in the ability of zebra mussels to selectively ingest high-quality organic particles when the organic content of the seston is low. Ingestion, assimilation efficiency, assimilation rate and scope for growth were all higher when seston was amended with an addition of a natural assemblage of algae. Food quality may be a better indicator of environmental conditions suitable for growth than food quantity. These results suggest that the conditions of high suspended inorganic sediment concentrations in large turbid rivers represent a difficult growth environment for the zebra mussel. Received: 12 May 1997 / Accepted: 7 July 1998     

Changes in the distribution and abundance of Dreissena polymorpha within lakes through time

Dreissena polymorpha population densities and biomass were followed in three Belarusian lakes with different trophic status over a 12-year period subsequent to initial colonization. In all three lakes zebra mussel population densities did not change once they reached a maximum. Application of the Ramcharan et al. [1992. Canadian Journal of Fisheries and Aquatic Sciences 49: 2611–2620] model for predicting population dynamics of zebra mussels was accurate for two of the three lakes studied. Population density appears to depend on the time since initial colonization, relative abundance of substrate available for colonization, lake morphometry and trophic type. Zebra mussel distribution within lakes was highly patchy, but the degree of dispersion decreased over time after initial colonization, which may be a result of saturation of suitable substrates by zebra mussels as populations increase and reach carrying capacity. In lakes where submerged macrophytes are the dominant substrate for zebra mussel attachment, populations may be less stable than in lakes with a variety of substrates, which will have a more balanced age distribution, and be less impacted by year to year variation in recruitment. Dreissena polymorpha usually reach maximum population density 7–12 years after initial introduction. However, the timing of initial introduction is often very difficult to determine. Both European and North American data suggest that zebra mussels reach maximum density in about 2–3 years after populations are large enough to be detected.     

Seasonal change in the spatial utilization of host mussels in relation to ovipositor length by female rosy bitterling Rhodeus ocellatus kurumeus

The seasonal changes in ovipositor length and utilization patterns of mussels for oviposition in the rosy bitterling Rhodeus ocellatus kurumeus (Cyprinidae) were investigated in a field experiment and field surveys during the breeding period (April to August). The mean length of ovipositors at oviposition was short at the start (early April) and end (July) of the breeding period. Females with long ovipositor at oviposition were collected between mid‐April and June. Mark‐and‐recapture data showed that ovipositor length at oviposition changed rhythmically throughout the breeding period, shortening and lengthening as the female entered the spawning and resting phase. The density of rosy bitterling embryos in mussels increased between April and June, peaking in May, but decreasing in July. The position of eggs on mussel gills varied from close to the exhalant siphon to deeper inside the gill during April, and periodically thereafter. There was a positive correlation between ovipositor length at oviposition and the distance from exhalant siphon of mussels to eggs deposited by females, suggesting that ovipositor length at oviposition determined the position of eggs deposited on a mussel gill. Because dissolved oxygen in mussel gills decreased with the density of bitterling embryos, suitable positions for embryo survival in gills changed with embryo density. By changing ovipositor length at oviposition, females might be able to spawn their eggs in a position that maximizes embryo survival. Thus, plasticity in ovipositor length at oviposition may play an important role as an adaptation of rosy bitterling in utilizing mussels when their quality as a spawning substratum fluctuates seasonally.     

An indirect effect of biological invasions: the effect of zebra mussel fouling on parasitisation of unionid mussels by bitterling fish

Invasive species represent a major threat with both direct and indirect effects on natural ecosystems, including effects on established and coevolved relationships. In a series of experiments, we examined how the interaction between two native species, a unionid mussel (Unio pictorum) and the European bitterling (Rhodeus amarus), a fish that parasitises unionids, was affected by the non-native zebra mussel (Dreissena polymorpha). The zebra mussel fouls hard substrates, including shells of living unionids, and its presence is often associated with a decrease in population density of native unionid mussels. Bitterling lay their eggs into live unionids and the embryos develop inside their gills. Using a range of zebra mussel densities, we demonstrated that zebra mussel fouling had a negative effect on the number of bitterling eggs inside the mussel host, with abundances of 5–10 zebra mussels (shell size 15–25 mm) per unionid critical for bitterling ability to utilise the host. In a further experiment, we found that bitterling did not discriminate between unfouled unionids and those fouled with five zebra mussels. Most ovipositions into fouled hosts, however, were unsuccessful as eggs failed to reach the unionid gills. We discuss implications of such unsuccessful ovipositions for bitterling recruitment and population dynamics.     

Predator‐induced morphological defences in two invasive dreissenid mussels: implications for species replacement

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The effect of seasonal variation in selective feeding by zebra mussels (Dreissena polymorpha) on phytoplankton community composition

1. To investigate the impact of zebra mussels ( Dreissena polymorpha ) on phytoplankton community composition, temporal variability in selective feeding by the mussels was determined from April to November 2005 in a natural lake using Delayed Fluorescence (DF) excitation spectroscopy.
2. Selective grazing by zebra mussels varied in relation to seasonal phytoplankton dynamics; mussels showed a consistent preference for cryptophytes and avoidance of chlorophytes and cyanobacteria. Diatoms, chrysophytes and dinoflagellates responded differentially to zebra mussel grazing depending on their size. Analysis of excreted products of the zebra mussels revealed that in addition to chlorophytes and cyanobacteria, phytoplankton >50  μ m and very small phytoplankton (≤7  μ m) were largely expelled in pseudofaeces.
3. The zebra mussel is a selective filter-feeder that alters its feeding behaviour in relation to phytoplankton composition to capture and ingest high quality phytoplankton, especially when phytoplankton occur in preferred size ranges. Flexibility of zebra mussel feeding behaviour and variation in susceptibility among phytoplankton groups to mussel ingestion indicate that invading zebra mussels could alter phytoplankton community composition of lakes and have important ecosystem consequences.     

Testing a typology system of running waters for conservation planning in Hungary

The zebra mussel (Dreissena polymorpha) and its congener the quagga mussel (Dreissena rostriformis bugensis) are both invaders in freshwater, but have very different invasion histories, with zebra mussels attaining substantially faster rates of spread at virtually all spatial scales. However, in waterbodies where they co-occur, D. r. bugensis can displace D. polymorpha. To determine if the mechanisms for this displacement are associated with different survival and growth, we kept mussels in flow-through tanks for 289 days with two temperature regimes that mimicked the natural surface water (littoral zone) and hypolimnion conditions of Lake Erie. For the littoral zone regime, we used water directly from the surface of Lake Erie (range 4–25°C, average 11.9 ± 0.6°C). For the profundal zone treatment, Lake Erie surface water was chilled to about 6°C (range 5–8°C, average 6.2 ± 0.6°C) for the full duration of the experiment. For each of these temperature regimes, we used three replicate tanks with only zebra mussels present and three replicate tanks with only quagga mussels (150 ind./tank each), and three replicate tanks with both species (75 ind./tank of each species). Quagga mussels had higher survivorship and grew more than zebra mussels in all treatments. For both species, the size of the mussel entering the winter was critical for survivorship. Larger mussels had a higher survival over the winter in all treatments. For both species, there was a survivorship and growth tradeoff. In the warmer littoral zone treatment both species had higher growth, but lower survival than in the colder profundal zone treatment. Surprisingly, although quagga mussels outperformed zebra mussels, zebra mussel survivorship was better when they were faced with competition by quagga mussels than with just intraspecific competition. In addition, quagga mussels suffered size-specific mortality during the growing season only when facing interspecific competition with zebra mussels. Further experiments are needed to determine the possible mechanisms for these interspecific effects.     

The zebra mussel (Dreissena polymorpha), a new pest in North America: reproductive mechanisms as possible targets of control strategies

Summary

The zebra mussel (Dreissena polymorpha) has spread rapidly in temperate fresh waters of North America since its introduction into the Great Lakes in 1985 or 1986. It attaches to hard substrates, forming layers, occluding water intakes, encrusting and killing native mussels, filtering algae in competition with other planktivores, and possibly interfering with fish spawning. It reproduces prolifically, suggesting that an approach to its control may be by controlling its reproduction. Previous literature suggests that spawning in bivalves is regulated by both environmental and internal chemical cues. A suggested sequence is that phytoplankton chemicals initially trigger spawning; chemicals associated with gametes provide a species-specific pheromonal positive feedback for spawning; and the response to environmental chemicals is mediated internally by serotonin (5-HT). The role of 5-HT in zebra mussels is under investigation. Both males and females can be induced to spawn by either injection or external application of 5-HT. The response can also be activated by 8-hydroxy-2-(di-n-propylamino)-tetralin, an agonist at 5-HT_1A receptors. HPLC analysis has detected 5-HT as the major biogenic amine in both male and female gonads. 5-HT immunocytochemistry demonstrates nerves containing serotonergic fibers innervating gonads of both males and females, with prominent varicosities surrounding the follicles in both sexes. A role of 5-HT in mediating spawning responses in zebra mussels is thus strongly supported. These studies have shown that reproductive behavior of zebra mussels can be modified by outside chemicals, a property that may be exploited for purposes of control.     

Phosphorus recycling by zebra mussels in relation to density and food resource availability

Using flow-through microcosms, we examined phosphorus (P) recycling by zebra mussels under conditions of nearly constant food resource supply and varying zebra mussel population densities (600–5200 ind./m²). At all density levels, zebra mussels filtered substantial algae, measured as chlorophyll biomass. Because chlorophyll biomass inputs were low throughout the study, zebra mussel biomass-specific rates of chlorophyll filtration declined with increasing density, suggesting food resource limitation at the higher densities. We observed net total P export and high zebra mussel biomass-specific rates of P recycling over time in microcosms at high zebra mussel densities. In systems with a low zebra mussel density, net total P export did not occur over time. Our results suggest the occurrence of P remineralization by zebra mussels and net loss associated with emaciation during periods of temporary starvation. These findings have implications for P dynamics since zebra mussels can be subjected to periods of starvation over seasonal and annual time scales.     

Annual spawning of the hydrothermal vent mussel, Bathymodiolus azoricus, under controlled aquarium, conditions at atmospheric pressure

Dense macrofaunal communities of modioliform mussels are a major component of many hydrothermal vent and cold seep ecosystems. The hydrothermal vent mussel Bathymodiolus azoricus, that dominates hydrothermal vent communities near the Azores Triple Junction, can be maintained in aquaria at atmospheric pressure. Cages containing these mussels were placed over diffuse vent outlets and recovered at different times. Cages recovered in January 2003 contained mussels with ripe gonads while those recovered in July, August and November 2001 and in April 2003 did not. Mussels collected post-spawning in April 2003 spawned in the aquaria in January 2004. Young mussels recruited to the cages in April 2003. The data support a main single period of spawning and of juvenile recruitment for B. azoricus.     

Putative serotonin reuptake inhibitor-induced spawning and parturition in freshwater bivalves is inhibited by mammalian 5-HT2 receptor antagonists

Selective serotonin reuptake inhibitors (SSRIs) can mimic the physiological actions of serotonin, and in bivalve molluscs they induce zebra mussel spawning and fingernail clam parturition. We have elucidated further the pharmacology of SSRI-induced spawning and part-urition by blocking these reproductive processes with two mammalian 5-HT(2) receptor antagonists, cyproheptadine and mianserin. These two antagonists were potent inhibitors of both spawning and parturition induced by the SSRIs fluvoxamine, fluoxetine, and zimelidine. In zebra mussels, both cyproheptadine and mianserin significantly blocked spawning induced by fluvoxamine and by zimelidine. In the fingernail clams Sphaerium spp., both cyproheptadine and mianserin blocked fluvoxamine-induced parturition. A possible mechanism of action for SSRI-induced spawning and parturition in bivalves is suggested.     

Factors affecting seasonal mortality of rosy bitterling (Rhodeus ocellatus kurumeus) embryos on the gills of their host mussel

I investigated the seasonal change in factors affecting embryonic mortality in the rosy bitterling, Rhodeus ocellatus kurumeus, a freshwater fish that spawns on the gills of living unionid mussels. Research was conducted in a small pond during 1999 and 2001 in which bitterling were provided with Anodonta sp. mussels for spawning. Bitterling spawned between April and July, peaking mid–late May. Seasonal survival rate of bitterling embryos in their mussel hosts was unimodal, with a peak between late April and mid May (about 70% of total spawnings). In mid April, survival was about 50%. The lowest survival was from late May to July (0%). Losses of bitterling embryos from mussels were identified by ejections from the mussel host. Ejections were categorized as either ejections of live embryos, or ejections of embryos that died in the mussel and were subsequently expelled from the mussel. Ejection rates of live embryos were higher in the earlier part of the spawning period (early–mid April) and dead embryo ejections in the later period (after June). The ejection rate of live embryos was higher among younger embryos earlier in the season, probably because of the incomplete development of morphological and behavioural traits associated with maintaining the embryo inside the mussel gill chambers, and as a consequence of a more protracted developmental period at low temperatures making them more susceptible to ejection. The ejection rate of dead embryos was higher in older embryos later in the season, and in larger mussels and at high embryo densities. The survival of embryos in mussels was probably related to oxygen availability, with mortalities probably caused by asphyxiation. Increased embryo mortalities may arise through competition among embryos, between embryos and mussel, and ambient dissolved oxygen levels. The optimal period for bitterling to spawn may represent a balance between two opposing factors; with positive and negative effects of a seasonal rise in temperature directly affecting embryonic growth rate and oxygen availability.An erratum to this article can be found at This revised version was published online January 2005 with the correction of the authors name.