Rhodeus Sericeus Research Articles

The action of bed-slope the fish community’s composition of the watercorses on the Zagyva-Tarna watersystem

Between 2003 and 2005 faunistic survey was completed in the sections of the low- and highland watercourses of the water-regime of the Zagyva and Tarna rivers. On the basis of bed-slope, dominance and frequency of the fish species the following sections can be divided in the examined streams:1./ Upper-chub-zone: the bed-slope is above 15 m/km. The typical species of this zone – the frequency of these is more then 50%, the dominance is more than 5% – is the Barbatula barbatula, but in the case of bigger streams Leuciscus cephalus is also characteristic.2./ Middle-chub-zone: the bed-slope is between 5-15 m/km. The typical fishspecies–besides the Barbatula barbatula and Leuciscus cephalus – is the Gobio gobio.3./ Down-chub-zone: in the case of larger watercourses (Zagyva and Tarna rivers) the bed-slope is 1,2-5 m/km, at smaller streamlets it is 1,7-5m/km. The typical fishspecies–besides the 3 above mentioned species–are the Cobitis elongatoides, the Alburnus alburnus and in the watersystem of Tarna the Alburnoides bipunctatus.4./ Top-perch-zone: the bed-slope is 0,3-1,2 m/km, or at smaller streamlets it is 0,3-1,7 m/km. The typical fishspecies–besidestheCobitis elongatoides, the Alburnus alburnus and the Alburnoides bipunctatus - are the Rutilus rutilus, the Gobio albipinnatus ant the Rhodeus sericeus.

Temporal and spatial distribution of glochidial larval stages of European unionid mussels (Mollusca: Unionidae) on host fishes

Glochidia are the larval stage of freshwater unionid mussels that parasitize the fins and gill apparatus of fish. A total of 22 fish species were examined for the presence of glochidia whose distribution on individual hosts was studied on three common fish species, the roach Rutilus rutilus (L.), perch Percafluviatilis L. and bitterling Rhodeus sericeus (Pallas). Between 1997 and 1999, the fish were obtained from the rivers Morava and Kyjovka and surrounding water pools in the Czech Republic. The glochidia of two genera, Unio and Anodonta, were found. Anodonta glochidia were observed on 10 fish species, Unio glochidia on 17 fish species. There was a difference in spatial distribution of glochidia on the body of the host fish. Unio glochidia were predominantly located on the gills, whereas most Anodonta glochidia were found on the fins, with the highest numbers of glochidia were observed on the margin of the pectoral fins. For the gill apparatus, Unio glochidia were found predominantly on the second and third arch. Anodonta glochidia were predominantly found during winter and spring (November-May), whereas Unio glochidia were more abundant during May and June. The number of glochidia was positively correlated with fish length in perch highly infected by Anodonta glochidia and perch infected by Unio glochidia. Of the three fish species, the highest occurrence of parasites was found on perch with fewer observed on roach. In spite of the close relationship between bitterling and unionid mussels, glochidiosis was rare on this fish species.

The costs and benefits in an unusual symbiosis: experimental evidence that bitterling fish (Rhodeus sericeus) are parasites of unionid mussels in Europe

Interspecific symbiotic relationships involve a complex network of interactions, and understanding their outcome requires quantification of the costs and benefits to both partners. We experimentally investigated the costs and benefits in the relationship between European bitterling fish (Rhodeus sericeus) and freshwater mussels that are used by R. sericeus for oviposition. This relationship has hitherto been thought mutualistic, on the premise that R. sericeus use mussels as foster parents of their embryos while mussels use R. sericeus as hosts for their larvae. We demonstrate that R. sericeus is a parasite of European mussels, because it (i) avoids the cost of infection by mussel larvae and (ii) imposes a direct cost on mussels. Our experiments also indicate a potential coevolutionary arms race between bitterling fishes and their mussel hosts; the outcome of this relationship may differ between Asia, the centre of distribution of bitterling fishes, and Europe where they have recently invaded.

Females solicit sneakers to improve fertilization success in the bitterling fish ( Rhodeus sericeus )

In most species there is greater variance in reproductive fitness among males than females. A consequence is the evolution of alternative male mating tactics, including sneaking behaviour whereby males attempt to mate with females courted by other males, which could undermine female choice. In bitterling (Rhodeus sericeus), fish that lay their eggs on the gills of living freshwater mussels, sneaking is common and successful. Here we show that female bitterling can significantly improve their fertility with an increase in the number of spawning partners. Females perform conspicuous behaviours associated with spawning more frequently close to sneakers, and spawn more eggs close to high-quality sneakers. This is the first time that females have been shown to engage in behaviour that increases the probability of sneaking, and raises the possibility of a sexual conflict over the number and source of ejaculates during spawning in this species.

Benefits and costs to mussels from ejecting bitterling embryos: a test of the evolutionary equilibrium hypothesis

Two major hypotheses of host–parasite interactions have been proposed to explain cases where hosts do not defend themselves against parasites. The evolutionary lag hypothesis suggests that there has been insufficient time for a host response to evolve, whereas the evolutionary equilibrium hypothesis proposes that host defence does not evolve because it carries costs that outweigh the benefits. We tested potential benefits and costs of host defence in an unusual interaction, between a freshwater fish, the European bitterling, Rhodeus sericeus, and live unionid mussels that are used as hosts for the fish’s eggs. We found a significant reduction in ventilation rate of mussels that were incubating bitterling embryos, which became more severe with an increasing number of embryos. We tested the hypothesis that the risk of ejecting a mussel’s own larvae while ejecting bitterling embryos has constrained the evolution of a host response. This predicts that brooding female mussels would retain more bitterling larvae than males or nonbrooding females. This prediction was not supported: brooding female mussels contained as many larvae per accessible gill as, and fewer in total than, males or nonbrooding females. In summary, based on the costs and benefits of ejection that we measured, we found no evidence in support of the evolutionary equilibrium hypothesis. However, other differences between mussel species, such as gill structure as measured in this study, ventilation rates and differences in the distance eggs are lodged into the gills, may contribute to differences in egg ejection rates.

Sexual selection for male dominance reduces opportunities for female mate choice in the European bitterling (Rhodeus sericeus)

Sexual selection involves two main mechanisms: intrasexual competition for mates and intersexual mate choice. We experimentally separated intrasexual (male-male interference competition) and intersexual (female choice) components of sexual selection in a freshwater fish, the European bitterling (Rhodeus sericeus). We compared the roles of multiple morphological and behavioural traits in male success in both components of sexual competition, and their relation to male reproductive success, measured as paternity of offspring. Body size was important for both female choice and male-male competition, though females also preferred males that courted more vigorously. However, dominant males often monopolized females regardless of female preference. Subordinate males were not excluded from reproduction and sired some offspring, possibly through sneaked ejaculations. Male dominance and a greater intensity of carotenoid-based red colouration in their iris were the best predictors of male reproductive success. The extent of red iris colouration and parasite load did not have significant effects on female choice, male dominance or male reproductive success. No effect of parasite load on the expression of red eye colouration was detected, though this may have been due to low parasite prevalence in males overall. In conclusion, we showed that even though larger body size was favoured in both intersexual and intrasexual selection, male-male interference competition reduced opportunities for female choice. Females, despite being choosy, had limited control over the paternity of their offspring. Our study highlights the need for reliable measures of male reproductive success in studies of sexual selection.

Seasonal occurrence and metrical variability of Gyrodactylus rhodei Žitňan 1964 (Monogenea, Gyrodactylidae)

The seasonal dynamics of Gyrodactylus rhodei, a monogenean ectoparasite of bitterling (Rhodeus sericeus), was studied from June 2000 to May 2001 in the Kyjovka River, Czech Republic. A negative relationship between prevalence and intensity of infection of G. rhodei and water temperature was found. Metrical variability of the hard parts of the parasite haptor was studied throughout the sampling season. A negative relationship between water temperature and the size of the hard parts of the G. rhodei haptor was evident in the measurements of the total length of the marginal hooks, the sickle length of marginal hooks, anchors, anchor point and root, the width of the ventral bar and the membrane processes. Sequences of the partial ITS (rDNA) of specimens collected during the cold and warm seasons were analysed. Sequences of all studied parasite specimens were identical and there was no evidence of intraspecific variability in the sequenced region.

The importance of species interactions in conservation: the endangered European bitterling Rhodeus sericeus and its freshwater mussel hosts

AbstractSpecies conservation may be complicated in symbiotic interactions. Sub‐lethal impacts of human activities on one species may be lethal to other species on which they depend. We investigated the impact of two aspects of eutrophication – changes in phytoplankton and oxygen levels – on the interaction between a freshwater fish that is endangered in several countries, the European bitterling, Rhodeus sericeus, and the unionid mussel hosts that they require for spawning. Reduced concentrations of algae and oxygen each led to a greater proportion of bitterling embryos being ejected prematurely from the mussel hosts. Reduced algal concentrations also led to reductions in mussel ventilation rates and to mussels spending less time with their valves open. Bitterling mortality may have been due to three causes: direct effects of environmental conditions on embryonic survival, active expulsion by mussels when stressed and through responses by mussels that created internal conditions that bitterling embryos could not survive. Our study shows that the sub‐lethal effects of pollution on one species can have lethal effects on another species with which it interacts. Such interactions need to be considered in conservation programmes.

Male-male interference competition decreases spawning rate in the European bitterling ( Rhodeus sericeus )

We investigated the consequences of male-male interference competition associated with alternative male mating tactics in a freshwater fish, the European bitterling (Rhodeus sericeus). Male bitterling defend territories around living mussels and attract females to lay their eggs in the gill cavities of mussels. We experimentally manipulated spawning-site abundance and male density at two spatial scales. We showed that the total number of eggs spawned by females was constrained by the number of mussels available for oviposition. The effect was mediated by behavioral interactions among competing males because of variation in the Operational Sex Ratio (OSR) in close proximity to a mussel and not by a direct limitation in mussel capacity to accommodate the eggs. Both total and local male densities affected spawning behavior, and interacted in their effect on female spawning rate. Territorial male aggression caused courtship interruptions that prolonged the time until successful spawning and increased with male density. However, territoriality broke down at the highest male density, with a consequent stabilizing effect on spawning rate.

The reproductive ecology of the European bitterling (Rhodeus sericeus)

AbstractThe European bitterling Rhodeus sericeus Pallas is a freshwater fish belonging to the Acheilognathinae, a subfamily that has an unusual spawning symbiosis with freshwater mussels. Female bitterling possess long ovipositors that they use to place their eggs onto the gills of a mussel through the mussels' exhalant siphon. Males fertilize the eggs by releasing sperm into the inhalant siphon of the mussel. The embryos develop inside the mussel for approximately a month, eventually leaving the mussel as actively swimming larvae. Because they use a discrete spawning site that can be readily manipulated they have recently been used in studies to understand oviposition choice with respect to mate and spawning site quality, host–parasite coevolution, alternative male mating tactics, and linking behavioural decisions with population dynamics. The current state of knowledge of bitterling reproductive ecology is reviewed and synthesized, and future directions for research on this species are proposed.

Molecular phylogenetic relationships of Eastern Asian Cyprinidae (pisces: cypriniformes) inferred from cytochrome b sequences.

Complete mitochondrial cytochrome b sequences of 54 species, including 18 newly sequenced, were analyzed to infer the phylogenetic relationships within the family Cyprinidae in East Asia. Phylogenetic trees were generated using various tree-building methods, including Neighbor-joining (NJ), Maximum Parsimony (MP) and Maximum Likelihood (ML) methods, with Myxocyprinus asiaticus (family Catostomidae) as the designated outgroup. The results from NJ and ML methods were mostly similar, supporting some existing subfamilies within Cyprinidae as monophyletic, such as Cultrinae, Xenocyprinae and Gobioninae (including Gobiobotinae). However, genera within the subfamily "Danioninae" did not form a monophyletic group. The subfamily Leuciscinae was divided into two unrelated groups: the "Leuciscinae" in East Asia forming as a monophyletic group together with Cultrinae and Xenocyprinae, while the Leucisciriae in Europe, Siberia, and North America as another monophyletic group. The monophyly of subfamily Cyprininae sensu Howes was supported by NJ and ML trees and is basal in the tree. The position of Acheilognathinae, a widely accepted monophyletic group represented by Rhodeus sericeus, was not resolved.

Operational sex ratio and resource defence as predictors of the mating system in European bitterling

Operational sex ratio (OSR), the ratio of sexually active males to fertilizable females in a population, plays a central role in the theory of mating systems by predicting that the intensity of male–male competition and the degree of sexual selection increases as the OSR becomes increasingly male biased. At high values of OSR, however, resource defence theory predicts the breakdown of territoriality and a shift towards scramble competition with a decrease in sexual selection. The direction that correlations between OSR and resource competition and variance in mating success will take depends on the biology of the species of interest. We investigated the effects of male population density and male‐biased operational sex ratio on male mating tactics shown by a freshwater fish, the European bitterling, Rhodeus sericeus. This species spawns inside living unioneid mussels. Large males defended territories, were aggressive towards conspecifics under equal sex ratios and monopolized pair spawnings with females. The mating tactic, however, changed at high male density where large males ceased to be territorial and instead competed with groups of smaller males to release sperm when females spawned. This change in male behaviour from pair to group spawning has two ramifications for sexual selection. The intensity of sexual selection and variance in male mating success decrease, and the form of sexual competition changes from resource‐ to sperm competition. Thus, the use of alternative mating tactics renders the OSR unable to predict the direction of resource competition and variance in male mating success at high densities.

Bitterling as models for studies of sperm competition

Game theory models of sperm competition predict that sperm expenditure should depend on the intensity of sperm competition. Specifically, there should be an increase in sperm expenditure when a male has one competitor, but a decrease with additional competitors. European bitterling, Rhodeus sericeus, are good models for testing this theory because sperm release by males is readily observable and territorial males regularly compete with variable numbers of sneakers. Females spawn in living freshwater mussels (Unionidae) and males release their sperm over the inhalent siphon both before and after egg laying. Our experiments confirmed that both territorial and sneaker males adjusted their sperm expenditure to the number of participating males, following the pattern predicted by game theory models. The pattern, however, was not always upheld: males increased their aggression level after a female had spawned and no longer adjusted their sperm expenditure to the number of sneaker males. Thus, adjusted sperm expenditure and mate guarding were two alternative responses to increased risk of sperm competition. Males adopted the tactic of sperm adjustment, when mate guarding and aggression against sneakers were costly in decreasing female willingness to spawn.

The European bitterling: a model for oviposition decision

For oviparous species, oviposition decisions can have significant fitness consequences. The European bitterling (Rhodeus sericeus) is a cyprinid fish that lays its eggs on the gills of freshwater mussels. Because bitterling use a discrete spawning site that can be readily manipulated, they are unusually amenable to field and laboratory studies aimed at understanding the adaptiveness and consequences of oviposition decisions. Here I present data demonstrating the adaptiveness of oviposition decisions by female bitterling, and link these decisions, using game‐theory models, with bitterling population dynamics. I explain the proximate cues used by females in making oviposition choices, and how these respond to environmental variables. I also show how the oviposition decisions of males relate to the risk of sperm competition in mussels, and propose that oviposition choices may represent an inter‐sexual conflict in bitterling. Finally, I present behavioural and genetic data from mesocosm experiments to show the implications of global and local male densities for female oviposition opportunities, and consider the consequences for bitterling population dynamics.

The bitterling–mussel interaction as a test case for co‐evolution

The European bitterling Rhodeus sericeus(Cyprinidae) spawns in the gills of freshwater mussels (Unionidae) and shows some obvious adaptations to this type of spawning, such as the development of an ovipositor. Furthermore, recent studies have shown that the fish avoid species of mussels that have a high likelihood of ejecting their eggs prematurely. This leads to the question of whether the interaction between bitterling and mussels could represent a case of co‐evolution, involving evolutionary responses by both species to selection imposed by the other. The evidence for and against co‐evolution is reviewed, incorporating new results from two sets of experiments designed to test for adaptive choices by bitterling according to the mussels’ sex and reproductive state, as well as a preliminary study of potential benefits for mussels from exposure to bitterling. Host preferences by bitterling, both among and within mussel species, may indeed have evolved in response to differences in benefits for offspring survival. There is no evidence yet for any benefits to mussels from receiving eggs, whereas there are costs due to reduced ventilation rates when the gills contain bitterling eggs. While there are differences among mussel species and individuals in their tendency to reject bitterling embryos, these differences do not provide strong evidence for co‐evolution. For example, they may reflect differences in host physiology such as ventilation rate and generalized responses to expelling objects from their gills. Therefore, while bitterling are well adapted for their obligate spawning relationship with mussels, it has been much more difficult to find evidence for adaptations by mussels for dealing with bitterling. This suggests that any co‐evolutionary dynamics between bitterling and mussels may be asymmetric, with stronger responses to selection by the fish than by mussels.

Operational sex ratio and alternative reproductive behaviours in the European bitterling, Rhodeus sericeus

We investigated the effects of male population density and male-biased operational sex ratio (OSR) with constant and limited resource density on male mating tactics shown by a freshwater fish, the European bitterling, Rhodeus sericeus. This species spawns inside living unionid mussels. Large males defended territories and were aggressive towards conspecifics under equal sex ratios. They also monopolised pair spawnings with females, releasing 98% of all sperm clouds during mating. However, the mating tactic changed at high male density where large males ceased to be territorial and instead competed with groups of smaller males to release sperm when females spawned. Large, medium and small males now obtained 61%, 33%, and 6% of sperm releases respectively, thereby reducing the opportunity for sexual selection by half. Females spawned at equal rates in the two densities of males, despite lower courtship at high density. These results run counter to the usual expectation that an increasingly male-biased OSR should lead to higher variance in male mating success. Instead, the use of alternative reproductive behaviours by males can lead to lower resource competition and mating variance at high male densities.

Isolation of microsatellite loci in the freshwater fish, the bitterling Rhodeus sericeus (Teleostei: Cyprinidae)

AbstractWe isolated 30 microsatellites from the freshwater fish, the bitterling Rhodeus sericeus. Twenty‐three microsatellite sequences possessed sufficient flanking DNA from which to design primers. Twelve loci were characterized and all were found to be polymorphic. These loci were isolated and characterized as part of a project to investigate the consequences of male alternative mating tactics and sperm competition using the bitterling as a model species.

Assessment of sperm competition by European bitterling, Rhodeus sericeus

We investigated male assessment of sperm competition in the bitterling, Rhodeus sericeus, a freshwater fish that spawns on the gills of living unionid mussels. Field experiments showed males increased their inspection rate of mussels into which a testis solution containing sperm had been experimentally released. Males avoided leading females to mussels that contained high numbers of embryos, but did not alter their leading behavior in response to the presence of sperm. In laboratory experiments males also increased their inspection rate of mussels into which a testis solution had been released and also failed to alter their leading behavior in response to the presence of sperm in mussels. However, males avoided leading females to mussels in close proximity to other males, and thereby may avoid sperm competition. In a second field study, territorial males were shown to ejaculate into mussels at a low rate in the absence of competitors, increase the frequency of ejaculations in competition with a rival, then decrease relative ejaculate expenditure as the number of competing males increased. Observed data were shown to be significantly correlated with predicted estimates of ejaculate expenditure for a model of sperm competition intensity. We discuss our results in the context of adaptive responses of males to sperm competition.

Parasite communities of freshwater fish under flood conditions.

The parasite communities of the freshwater fish species roach ( Rutilus rutilus), bitterling ( Rhodeus sericeus amarus) and perch ( Perca fluviatilis) were investigated during a major flood. Differences in parasite community structure due to changes in the host environment were expected. We therefore tested for differences in parasite species abundance and diversity as a consequence of the flood. Potential changes in parasite community structure due to seasonal influences were also considered. We found differences in the composition of parasite species and the proportion of ecto- and endoparasites in the three host species. The parasite community of roach was richer in species than those of perch and bitterling. Roach harboured more ectoparasite than endoparasite species. Parasite communities of both perch and bitterling were richer in endoparasite species and had a lower number of ectoparasite species. Parasite communities of the three different host species responded in different ways to environmental change. Considering all parasite species, no effect of either season or flood was found on the species diversity in either perch or bitterling. The flood caused an increase in the parasite species diversity in roach. For roach and perch, higher parasite abundance were found before the flood compared to after flood. However, no difference was found between the different seasons after the flood. By contrast, bitterling had the highest parasite abundance in the season after the flood. When analysing ecto- and endoparasites separately, a decrease in ectoparasite abundance was found in roach and perch, probably related to the flood, and due mainly to a decrease in monogenean species. A significant increase in the abundance of endoparasites was found in perch. In bitterling, the post-flood increase in ectoparasites was due to a high abundance of Gyrodactylus spp. We conclude that the parasite communities of the three fish species responded in different ways to the flood. This may be due to changes in the host life history strategy or in the immune response in conditions of stress.

Adjustments of ejaculation rates in response to risk of sperm competition in a fish, the bitterling (Rhodeus sericeus).

Game theory models of sperm competition predict that within species, males should increase their sperm expenditure when they have one competitor, but decrease expenditure with increasing numbers of competitors. So far, there have been few tests or support for this prediction. Here, we show that males of a freshwater fish, the European bitterling, Rhodeus sericeus, do indeed adjust their ejaculation rate to the number of male competitors by first increasing and then decreasing their ejaculation rates as the number of competitors increases. However, this occurred only under restricted conditions. Specifically, the prediction was upheld as long as no female had deposited eggs in the live mussels that are used as spawning sites. After one or more females had spawned, males did not decrease their ejaculation rates with the number of competitors, but instead they became more aggressive. This indicates that decreased ejaculation rate and increased aggression are alternative responses to increased risk of sperm competition.