Reciprocal Effects between Burying Behavior of a Larval Dragonfly (Odonata: Macromia illinoiensis) and Zebra Mussel Colonization

Invasive zebra mussels (Dreissena polymorpha) often colonize dragonfly larvae, especially spawling species whose survivorship to emergence as terrestrial predators is consequently reduced. Using individuals of the sprawler, Macromia illinoiensis, as their own controls, we compared the burying behavior of penultimate instar larvae before (i.e. baseline) and after their colonization by zebra mussels under ambient conditions. Individuals that took longer to bury themselves when mussel-free had a higher rate of colonization by mussels over a five-day period compared to those that buried faster. In contrast, the depth at which individuals buried when mussel-free was not predictive of subsequent colonization rate. Although mean bury time did not differ between baseline and when an individual carried one or more mussels, colonized larvae buried more shallowly than when mussel-free. Moreover, attached mussels increased the risk of subsequent colonization by zebra mussels. After naturally losing all of their attached mussels, bury time and depth of individuals did not differ from their baseline behavior, indicating that the changes in the behavior of colonized individuals were due to mussel loads and not their time in captivity. Under natural conditions, the positive feed-back between mussel attachment and increasing vulnerability to colonization helps explain how mussel loads, which are lost at molting, can accumulate quickly over the duration of the final larval stadium. Because zebra mussel attachment decreases the crypsis that that a M. illinoiensis gains from burying, the invasive mussel may also make dragonfly larvae more detectable to visual predators.     

Effects of zebra mussel attachment on the foraging behaviour of a larval dragonfly,Macromia illinoiensis

1. Larvae of Macromia illinoiensis Walsh are often colonised by the zebra mussel, Dreissena polymorpha Pallas, a recent invader to North America. To determine how mussel attachment affects an individual's foraging behaviour, we quantified capture of Hexagenia limbata Hexes mayfly prey and the distance moved by newly‐molted final instars before and after an individual's colonisation with zebra mussels. 2. In night trials, larvae sprawled above the sand, and caught more mayflies than individuals in daytime trials, but the estimated distance travelled did not differ. When resting under a layer of sand with only its eyes exposed during the day, an individual could capture a mayfly prey using a sit‐and‐wait ambush strategy. When sprawled above the sand, some larvae caught prey that rested on their legs. 3. When mussel‐free, individuals captured more prey than they did when carrying zebra mussels, although mussel attachment per se did not affect the estimated distance that a larva moved. 4. During day trials, but not night ones, the increasing mussel load of colonised individuals decreased prey capture and the distance moved in an apparent step‐wise function. Although the number of mussels carried did not differ, night foragers carried a heavier load. Independent of time of the day, the distance an individual travelled when mussel‐free was predictive of the number of prey it caught when colonised, suggesting that the greater general activity of some individuals helped mitigate negative effects that mussel attachment had on prey capture. 5. Our results add to a growing number of negative effects of zebra mussel colonisation on sprawling and hiding dragonfly larvae. Although the impact of these costs on dragonfly populations remains to be determined, a decrease in this guild of predators whose life cycle spans aquatic and terrestrial habitats might have cascading effects across ecosystems.     

Indigenous microflora and opportunistic pathogens of the freshwater zebra mussel, Dreissena polymorpha

Freshwater fouling invertebrate zebra mussels (Dreissena polymorpha) harbor a diverse population of microorganisms in the Great Lakes of North America. Among the indigenous microorganisms, selective species are opportunistic pathogens to zebra mussels. Pathogenicity to zebra mussels by opportunistic bacteria isolated from the mussels was investigated in this study. Among the more than 30 bacteria isolated from temperature-stressed mussels, Aeromonas media, A. veronii, A. salmonicida subsp. salmonicida, and Shewanella putrefaciens are virulent pathogens to juvenile zebra mussels. Inoculation of a bacterial concentration of A. media, A. salmonicida subsp. salmonicida and S. putrefaciens at 10⁷ cells per zebra mussel resulted in 100% mortality within 5 days, and only 64.9% for A. veronii. In contrast, mortality was less than 12.3% following inoculation of a sterile phosphate buffer solution as a control. In addition, mortality was dependent on the size of the pathogen population used in inoculation and the incubation temperature, indicating the close relationship between the bacterial population and subsequent death. On the mussel tissue, a dense microbial population was evident from the moribund mussels viewed with Scanning Electron Microscope (SEM). Opportunistic bacteria invaded and destroyed the D. polymorpha tissue after 7 days of incubation when the bacterial inoculation was larger than 10⁵ per zebra mussel. Our results suggest that mussels are reservoirs of opportunistic pathogenic microorganisms to aquatic organisms and humans and a better understanding of the microbial ecology of the mussels will provide insights to the possible health hazards from these microorganisms.     

Epiphytic Refugium: Are Two Species of Invading Freshwater Bivalves Partitioning Spatial Resources?

Enumeration of benthic (bottom dwelling) and epiphytic (attached to plants) zebra and quagga mussels (Dreissena polymorpha and D. bugensis, respectively) at Lake Erie near-shore sites in fall of 2000 revealed an unexpected prevalence of the zebra mussel on submerged plants. Even at Buffalo, New York, USA, where benthic dreissenids have been 92–100% quagga mussel since 1996, zebra mussels constituted 30–61% of epiphytes numerically. This may reflect a partitioning of settling space consistent with interspecific competition. A seasonal epiphytic refugium might allow the zebra mussel to persist even where the benthos is almost exclusively quagga mussel. This revised version was published online in July 2006 with corrections to the Cover Date.     

Onshore–offshore variations of copepod community in northern South China Sea

Previous studies which have tested the feeding preferences of shredders for fungal species and the food quality of fungi used detritus uniformly colonized by a fungus, which is not the case for decaying leaves in streams. It is not known whether shredders in different development stages exhibit variations in feeding preference and larval performance. This study examined the feeding preferences and the growth of the third and the fifth instars of Pycnopsyche gentilis larvae using fungal-colonized patches and whole leaves, respectively, having different fungal species compositions (Alatospora acuminata, Anguillospora filiformis, Articulospora tetracladia, Tetrachaetum elegans, and all species combined). The aquatic hyphomycetes used were co-dominant on leaves in the stream inhabited by the caddisfly. During 14 d of feeding, the larvae of both instars did not show significant differences in feeding preferences for the patches growing on oak leaves, although the third instar larvae were slightly more selective than the fifth instar larvae. When fed with maple leaves for 18 d, larval growth rates, gross growth efficiencies, and survivorship were not significantly different among the fungal treatments. However, the larval growth of both instars fed with fungal-colonized leaves was always significantly greater than the growth of larvae fed with diets of uncolonized leaves. The third instar larvae grew faster than the fifth instar larvae, but the growth efficiencies of the two instars were similar. These results suggest that P. gentilis larvae exhibit less selectivity in their feeding than other caddisfly shredders that have been examined and that the dominant fungi colonizing leaves in their habitat are similar in palatability and food quality for this shredder. Handling editor: B. Oertli     

Invasive zebra mussel colonisation of invasive crayfish: a case study

We investigated the interaction between two invasive invertebrate species in a shallow Central European flooded sandpit: the epibiosis of Ponto-Caspian zebra mussels Dreissena polymorpha on the American crayfish Orconectes limosus. Between 2004 and 2005, we followed the seasonal variation in number and size of the mussels attached to crayfish bodies, and microhabitats preferred by mussels. The proportion of crayfish colonised by mussels varied seasonally: in spring and early summer it was consistently over 75%, afterwards it dropped temporarily due to loss of bivalves during the crayfish moult, and later increased again due to re-colonisation by often relatively large juvenile mussels. Three different pathways of mussel settlement on crayfish hosts are likely: (1) primary settlement of free-swimming pediveliger larvae; (2) secondary settlement of plantigrade mussels and juveniles; (3) active re-attachment of grown mussels from the substrate to crayfish. This epibiosis was promoted by lack of suitable substrates at the studied locality. Electronic supplementary material The online version of this article (doi: ) contains supplementary material, which is available to authorized users.     

Comparative susceptibility of larval instars and pupae of the western corn rootworm to infection by three entomopathogenic nematodes

As a first step towards the development of an ecologically rational control strategy against western corn rootworm (WCR; Diabrotica virgifera virgifera LeConte, Coleoptera: Chrysomelidae) in Europe, we compared the susceptibility of the soil living larvae and pupae of this maize pest to infection by three entomopathogenic nematode (EPN) species. In laboratory assays using sand-filled trays, Heterorhabditis bacteriophora Poinar and H. megidis Poinar, Jackson & Klein (both Rhabditida: Heterorhabditidae) caused comparable mortality among all three larval instars and pupae of D. v. virgifera. In soil-filled trays, H. bacteriophora was slightly more effective against third larval instars and pupae, and H. megidis against third larval instars, compared to other developmental stages. In both sand and soil, Steinernema feltiae (Filipjev) (Rh.: Steinernematidae) was least effective against second instars. In conclusion, all larval instars of D. v. virgifera show susceptibility to infection by all three nematodes tested. It is predicted that early application against young larval instars would be most effective at preventing root feeding damage by D. v. virgifera. Applications of nematodes just before or during the time period when third instars are predominant in the field are likely to increase control efficacy. According to our laboratory assays, H. bacteriophora and H. megidis appear to be the most promising candidates for testing in the field. I. Hiltpold similarly contributed to this paper as the first author.     

Proteomic approach to probe for larval proteins of the mussel Mytilus galloprovincialis

A proteomic approach was used to search for larval proteins specific to the mussel Mytilus galloprovincialis from Galicia in northwest Spain. The study included both a comparative analysis, through two-dimensional electrophoresis, of protein expression maps of the larvae of the mussel and of 5 abundant and commercially important bivalve species from the region (Ostrea edulis, Cerastoderma edule, Pecten maximus, Tapes decussatus, and Venarupis pullastra) and subsequent mass spectrometric analysis of some of the protein spots. A total of 18 spots were selected and isolated from gels of M. galloprovincialis larvae. From their relative position on the electrophoresis gels, 6 of these were clearly exclusive to the mussel species. However, it was not clear whether the other spots were shared by other species. To overcome this ambiguity, first an analysis using matrix assisted laser desorption ionization with time-of-flight (MALDI-TOF) was conducted on the 6 spots of Mytilus that could possibly be shared with only one species. The peptide mass fingerprinting was completely different for the proteins compared. This result confirmed that the 6 proteins were exclusively mussel proteins, but demonstrated the utility of this approach when working with species that are poorly represented at the protein level in databases.     

Occurrence of Conchophthirus acuminatus (Protista: Ciliophora) in Dreissena polymorpha (Mollusca: Bivalvia) along the River Shannon, Ireland

Samples of Dreissena polymorpha were collected at several sites along the River Shannon navigation in Ireland, to determine the occurrence and distribution of their obligate host-specific commensalistic ciliate, Conchophthirus acuminatus, in this newly invaded region. Mussels collected by various methods were fixed immediately in 75% ethanol, in which they were later dissected under a stereoscopic microscope, beginning with thorough flushing of the mantle cavity and removal of the gills. One ml of sediment flushed from the mantle cavity and dissection residue of each mussel was examined under a compound light microscope using brightfield, phase-contrast, and differential-interference-contrast optics. Of 180 mussels examined, 125 (69.44%) harbored C. acuminatus. The ciliates were invariably well fixed and easily identifiable in all preparations. Mean sampling intensity for infected mussels was 8.47 ciliates per ml of sediment. Both prevalence and sampling intensity varied between sites, but no pattern was discernible. The present results are consistent with other reports of C. acuminatus being the most widespread and abundant symbiont of D. polymorpha throughout Europe, often occurring where no other symbionts occur. Its occurrence in Ireland indicates introduction of the mussels as adults, since planktonic veliger larvae are not known to harbor ciliates. Following similar reasoning, it is possible that the earlier North American invasion by D. polymorpha included only veligers, since C. acuminatus has not been found on that continent. Using these simple and quick methods, the ciliates could be easily identified and counted to give general comparative data among sites regarding intensity and prevalence. Thus, this method has promise for future efforts to obtain basic information rapidly in newly invaded systems.     

Mate recognition inCryptomyzus aphids: copulation and insemination

The acceptability of three widely distributed Australian Menispermaceae,Tinospora smilacina Benth.,Sarcopetalum harveyanum F. Muell. andStephania japonica (Thunb.) Miers, as food for larvae of the fruitpiercing moth,Othreis fullonia (Clerck), was examined in three laboratory experiments. When larvae were presented with plant species individually total development times were shortest onT. smilacina and longest onS. japonica, despite relatively similar consumption rates within most instars.T. smilacina elicited greater (P<0.05) relative growth rates thanS. japonica in all instars except the 6th. In the second experiment, when larvae were allowed to select from each of the 3 plants, noS. japonica was chosen by 1 st instars and it represented only 3.7% of food consumed by 2nd instars. Significantly moreT. smilacina was eaten in each instar thanS. japonica, and more thanS. harveyanum except in the 2nd and 4th instars. The final experiment examined the abilities of larvae to switch hosts when forced after the 1st and 3rd instars. After the first or second food change largest average headcapsule widths were associated with feeding onT. smilacina as the most recent food. Feeding by final instars onT. smilacina also resulted in the shortest development time and highest puparial weights. While some larvae survived irrespective of plant sequence 83.3% of the recorded mortality occurred while larvae were exposed toS. japonica, principally during the 1st instar. These experiments lend support to field observations which suggest thatT. smilacina is a major host ofO. fullonia whileS. japonica is notS. harveyanum is probably an important alternate host whenT. smilacina is scarce.     

Invasion ecology of quagga mussels (<Emphasis Type="Italic">Dreissena rostriformis bugensis</Emphasis>): a review of evolutionary and phylogenetic impacts

Two invasive freshwater mussels, Dreissena rostriformis bugensis (quagga mussel) and D. polymorpha (zebra mussel), reveal differences in patterns and timing of their invasions in Europe. They belong to different clades in Dreissena phylogenetics: D. rostriformis bugensis genetically is coupled with the brackish water, lacustrine D. r. distincta and the two are believed to represent a single species. As such, the guaqqa mussel has environmental requirements that differ from the congeneric D. polymorpha. D. rostriformis bugensis invasions were confined to reservoirs of the Dnieper, Don and Dniester Rivers of the Black Sea basin. We recorded D. r. bugensis outside the Black Sea basin for the first time between 1992 and 2001, along the Volga River reservoir cascade including the Northern Caspian Sea shallows. This represents a 40-year invasion time lag since an invasion corridor through the Volgo-Don Waterway was established in 1952 (a corridor used extensively by many invertebrate species from the Black Sea region). We attribute the postponed invasion of Europe by D. r. bugensis, including peculiarities in establishment and its absence in fossil records, to its phylogenetically close relationship with D. r. distincta and its recent evolutionary origin. The relatively rapid range expansion of D. r. bugensis in eastern Europe during the past several decades was facilitated by human-mediated ecosystem transformation, notably impoundment of large eastern European rivers, that have allowed this species to utilize newly transformed ecosystems.     

Influence of toxic and non-toxic phytoplankton on feeding and survival of Dreissena polymorpha (Pallas) larvae

Grazing and survival of larvae of the zebra mussel, Dreissena polymorpha, on a green alga and cyanobacteria were studied in laboratory experiments. Clearance rates of the larvae were determined for Chlamydomonas reinhardtii (green alga), two non-toxic and two toxic Microcystis aeruginosa strains (Cyanobacteria). Clearance rates of larvae on non-toxic Microcystis were significantly higher than on toxic Microcystis. The clearance rate on Chlamydomonas reinhardtii was in between the clearance rates on toxic and non-toxic Microcystis strains and not significantly different from them. Effects of toxicity of Microcystis on the survival of zebra mussel larvae was investigated in a short-term experiment. Survival of larvae fed toxic Microcystis was lower than that of larvae fed non-toxic Microcystis, but higher than that of starved larvae. This may imply that, for survival of zebra mussel larvae, it is better to have bad quality (toxic) food than no food.     

环境DNA技术在贻贝入侵监测中的应用挑战

沼蛤(Limnoperna fortunei)和斑马贻贝(Dreissena polymorpha)是淡水系统中常见的入侵贻贝物种,对其种群规模的持续监测是入侵贻贝防治管控中的关键环节。随着分子生物学技术的发展,入侵物种监测中逐渐尝试利用环境DNA(eDNA)技术实现快速、灵敏检测。然而,在入侵物种引入-定植-扩散过程的监测中,eDNA技术的灵敏度及定量效果受到诸多因素的影响,给实际应用带来挑战。系统梳理了国内外学者利用eDNA技术监测沼蛤、斑马贻贝等入侵物种的研究进展;分析了eDNA技术的采样方案、引物设计、定量分析、质量保证、原位便携仪器设计等影响监测效率与准确率的关键环节;进一步探讨了eDNA技术在贻贝入侵监测中的优势和局限性,以及未来的改进方向。     

Invasive zebra mussel (Dreissena polymorpha) threatens an exceptionally large population of the depressed river mussel (Pseudanodonta complanata) in a postglacial lake

1. Freshwater mussels are in decline worldwide, with the depressed river mussel Pseudanodonta complanata being one of the rarest and most endangered species in Europe. Invasive mussels are suspected to be an important factor of decline, but there is little information on their interaction with native species.
2. This study analyzed densities, depth distribution, and individual sizes and weights in one of the largest known populations of P. complanata in Europe in relation to the co‐occurring invasive zebra mussel Dreissena polymorpha and other mussel species, using a systematic transect analysis.
3. Pseudanodonta complanata was the dominant unionid species in Lake Siecino reaching densities of up to 26 ind/m², with half of the specimens found at a water depth of 2.0–4.0 m. Densities were highest on sandy substrates in areas of underwater currents. In contrast, 67% of native Unio tumidus were found at depths < 1 m, indicating different habitat preference.
4. In the study area, 91% of P. complanata, 92% of U. tumidus, and all Anodonta individuals were fouled by D. polymorpha. The dreissenid:unionid mass ratio (mean ± SD; maximum) was 0.43 ± 0.56; 4.22 and 0.86 ± 1.87; 8.76 in P. complanata and U. tumidus, respectively. Pseudanodonta complanata fouled with D. polymorpha were impaired in their anchoring capability and had shell deformations potentially affecting shell closing and filtration activity. Fouling intensity was negatively correlated with unionid density, potentially leading to accelerated population declines.
5. The observed adverse effects of invasive zebra mussels on the depressed river mussel and the difficulties in eradicating established populations of invasive mussels suggest that D. polymorpha should be considered a serious threat to P. complanata. Therefore, the further spread of zebra mussels into habitats with native unionids needs to be avoided by all means.

     

Interactions among coexisting larval Odonata: an in situ experiment using small enclosures

Field experiments using small replicated enclosures focused on interactions between larval populations of Epitheca cynosura and Ladona deplanata (Odonata: Anisoptera) — two species that emerge in early spring. The presence of Epitheca reduced the total biomass of Ladona, but Ladona had no significant effect on Epitheca. These early-emerging species reduced the biomass of small instars of late-emerging Anisoptera which colonized enclosures during the experiments; and the late-emerging Anisoptera seem to have inhibited colonization by Zygoptera larvae. Results are consistent with the importance of predatory (cannibalism or mutual predation) interactions in this community.     

Antipredatory Activity of the Weevil Oxyops vitiosa: A Biological Control Agent of Melaleuca quinquenervia

The larvae of the leaf-feeding weevil Oxyops vitiosa, a biological control agent of Melaleuca quinquenervia, are covered with a viscous orange coating that is thought to protect against generalist predators. This coating is gradually lost as the larvae drop to the ground and pupate in subterranean pupal cells. To test the antipredator activity of this species, four immature life stages (early instars, late instars, prepupae, pupae) were exposed to a common generalist predator, the red imported fire ant Solenopsis invicta. Choice tests were conducted by placing an O. vitiosa individual and a control larva of the weevil Neochetina eichhorniae into an arena containing a S. invicta colony and observing subsequent ant behaviors. S. invicta workers contacted O. vitiosa early instars, late instars, and prepupae less frequently than control N. eichhorniae larvae, and upon contact S. invicta was less likely to behave aggressively toward these O. vitiosa life stages than toward N. eichhorniae larvae. However, S. invicta contacted, attacked, and consumed naked (nonencased) O. vitiosa pupae and N. eichhorniae larvae with equal frequency. Encased O. vitiosa pupae buried in sand were not attacked compared to susceptible encased pupae on the sand surface. By shifting from a chemical defense during the larval stages to a physical defense during the pupal stage, O. vitiosa reduces the risk of attack by this generalist predator.     

Guild structure in water mites (Unionicola spp.) inhabiting freshwater mussels: choice,competitive exclusion and sex

Summary Unionicolid water mites inhabit freshwater unionid mussels during the nymphal and adult stages of their life-cycle. Regular sampling of mussels from two sites in St. Mark's River, Fl. established that each of four species of water mite (Unionicola abnormipes, U. fossulata, U. serrata and U. formosa) occurred mainly in one or two of the mussel species available at each site.The role of preference for particular mussel species during host location was assessed for the first three mite species by choice experiments, in which mites were offered different mussel species simultaneously. In five out of six experiments, mites entered normally unused mussels as often as they did normally used ones. Additionally, a sexual difference in choice was found for U. fossulata, with males preferring one mussel species and females showing no preference. One mussel species, (Anodonta imbecilis), normally unused but chosen by mite species during the lab. experiments, is inhabited exclusively by the fourth mite species, U. formosa, in the field. An experiment showed that U. formosa excludes other mite species aggressively from Anodonta imbecilis.The results illustrate the sometimes misleading nature of simple sampling data as an indication of host specificity or host preference in parasites. They suggest also that the population dynamics of some parasites might be more fruitfully compared to unrelated, free-living species than to other parasites.     

The association between the phytoplankton, Rhopalosolen species (Chlorophyta; Chlorophyceae), and Anopheles gambiae sensu lato (Diptera: Culicidae) larval abundance in western Kenya

Algae are important food resources of the larvae of the African malaria vectors, Anopheles gambiae Giles and Anopheles arabiensis Patton (Anopheles gambiae sensu lato), and other zooplankton, but empirical evidence remains meager about the agal flora in ephemeral water bodies. The animals present in natural aquatic habitats in western Kenya were sampled from July to November 2002 to study abiotic and biotic environmental factors determining A. gambiae sl larval abundance. The five highest concentrations of third and fourth instars and pupae (hereafter referred to as old-stage larvae) were sampled in conjunction with the unicellular epizoic green algae, Rhopalosolen species (Chlorophyta; Chlorophyceae). Canonical correspondence analysis revealed that the presence of Rhopalosolen species was the most important determinant of the animal assemblage. The density of old-stage A. gambiae sl larvae was positively correlated with the presence of Rhopalosolen species, but the density of first and second instars of A. gambiae sl was not. The water bodies with Rhopalosolen sp. yielded larger mosquitoes in spite of the higher density of larvae. We demonstrated that the productivity of water bodies in terms of the larvae of malaria vectors can differ in magnitude depending on the agal flora. We discuss phytoplankton as a regulator of mosquito larval populations.     

Regulation ofHelicoverpa zea larval behavior by the parasitoidEucelatoria bryani

The parasitoidEucelatoria bryani Sabrosky regulates the larval behavior of its hostHelicoverpa zea (Boddie). Parasitized third, fourth and fifth instars burrow into the soil 0.7–3.4 days earlier than unparasitized larvae that normally enter the soil to pupate at the end of the fifth and final larval instar. Parasitized third instars molt once then burrow as fourth instars, one instar earlier than normal. WhenE. bryani pupariated on the soil surface in the field, none survived to the adult stage. However,E. bryani adults emerged from 49.2% of hosts that had burrowed into the soil. By accelerating the timing ofH. zea burrowing behavior and causing host larvae to enter the soil before death,E. bryani ensures its pupariation in an environment with improved protection against natural enemies and lethal temperatures.     

ENDOSYMBIONTS OF DREISSENA POLYMORPHA IN IRELAND: EVIDENCE FOR THE INTRODUCTION OF ADULT MUSSELS

Although zebra mussels (Dreissena polymorpha) have invaded watersacross Europe for over 200 years, they colonized Ireland onlywithin the past decade. To test the hypothesis that Irelandwas colonized by adult D. polymorpha, we examined mussels fromdifferent sites along the Lower Shannon River system in Irelandfor the presence of host specific and generalist endosymbionts.Withinthe mantle cavity and/or associated with zebra mussel tissueswe found species specific-ciliates (Conchophthirus acuminatusand Ophryoglena hemophaga) and generalist symbionts (the ciliateAncistrumina limnica, nematodes, oligochaetes and chironomids).We found a significant difference in the prevalence of symbiontsamong sites, but all mussels at all sites harboured one specialistspecies C. acuminatus, and all of the mussels at three of thefour sites also had the second specialist, O. hemophaga. Thus,with the introduction of D. polymorpha into Ireland, at leasttwo additional species, their host-specific symbionts C. acuminatusand O. hemophaga, have also been introduced. The presence ofthese symbionts in Ireland supports the hypothesis that adultzebra mussels were introduced into Ireland, rather than larvalstages. This contrasts with the introduction of zebra musselsto North America, where adult zebra mussels are devoid of host-specificsymbionts. (Received 8 June 2005; accepted 7 November 2005)