Low Susceptibility of Invasive Red Lionfish (Pterois volitans) to a Generalist Ectoparasite in Both Its Introduced and Native Ranges

Escape from parasites in their native range is one of many mechanisms that can contribute to the success of an invasive species. Gnathiid isopods are blood-feeding ectoparasites that infest a wide range of fish hosts, mostly in coral reef habitats. They are ecologically similar to terrestrial ticks, with the ability to transmit blood-borne parasites and cause damage or even death to heavily infected hosts. Therefore, being highly resistant or highly susceptible to gnathiids can have significant fitness consequences for reef-associated fishes. Indo-Pacific red lionfish (Pterois volitans) have invaded coastal habitats of the western tropical and subtropical Atlantic and Caribbean regions. We assessed the susceptibility of red lionfish to parasitic gnathiid isopods in both their native Pacific and introduced Atlantic ranges via experimental field studies during which lionfish and other, ecologically-similar reef fishes were caged and exposed to gnathiid infestation on shallow coral reefs. Lionfish in both ranges had very few gnathiids when compared with other species, suggesting that lionfish are not highly susceptible to infestation by generalist ectoparasitic gnathiids. While this pattern implies that release from gnathiid infestation is unlikely to contribute to the success of lionfish as invaders, it does suggest that in environments with high gnathiid densities, lionfish may have an advantage over species that are more susceptible to gnathiids. Also, because lionfish are not completely resistant to gnathiids, our results suggest that lionfish could possibly have transported blood parasites between their native Pacific and invaded Atlantic ranges.     

A universal method for the detection and identification of Aphidiinae parasitoids within their aphid hosts

Molecular methods are increasingly used to detect and identify parasites in their hosts. However, existing methods are generally not appropriate for studying complex host-parasite interactions because they require prior knowledge of species composition. DNA barcoding is a molecular method that allows identifying species using DNA sequences as an identification key. We used DNA amplification with primers common to aphid parasitoids and sequencing of the amplified fragment to detect and identify parasitoids in their hosts, without prior knowledge on the species potentially present. To implement this approach, we developed a method based on 16S rRNA mitochondrial gene and LWRh nuclear gene. First, we designed two primer pairs specific to Aphidiinae (Hymenoptera), the main group of aphid parasitoids. Second, we tested whether the amplified regions could correctly identify Aphidiinae species and found that 61 species were accurately identified of 75 tested. We then determined the ability of each primer pair to detect immature parasitoids inside their aphid host. Detection was earlier for 16S than for LWRh, with parasitoids detected, respectively, 24 and 48 h after egg injection. Finally, we applied this method to assess parasitism rate in field populations of several aphid species. The interest of this tool for analysing aphid-parasitoid food webs is discussed.     

Cleaner fish, Labroides dimidiatus, diet preferences for different types of mucus and parasitic gnathiid isopods

Cleaner fish, Labroides dimidiatus, prefer the mucus of the parrotfish, Chlorurus sordidus, to parasitic gnathiid isopods, the main items in their diet, indicating a major conflict between clients and cleaners over what the latter should eat during interactions. We tested whether the conflict varied with client species (and the quality of its mucus) and with the presence of blood in the gnathiids. First, we offered cleaners the choice between mucus of the parrotfish and that of the snapper, Lutjanus fulviflamma. When offered equal amounts of mucus on Plexiglas plates, cleaners readily developed a significant preference for the parrotfish mucus. Reducing the amount of parrotfish mucus by 75% made the preference disappear. In a second test, we offered the cleaners gnathiids that were or were not engorged with client fish blood. Cleaners showed no significant preference for either food item. Our results suggest that the degree of conflict between cleaners and clients may vary between species, depending on whether the latter have a preferred mucus. In contrast, the cleaners' lack of preference for engorged gnathiids benefits clients because it means that cleaners do not hesitate to eat unengorged gnathiids before the gnathiids harm the fish by removing blood or by transmitting blood parasites.     

Diurnal variation in the abundance of juvenile parasitic gnathiid isopods on coral reef fish: implications for parasite-cleaner fish interactions

 The dynamics of parasitic gnathiid isopod infestation on the fish Hemigymnus melapterus were examined at Heron Island, Great Barrier Reef, by measuring the abundance and feeding state of gnathiids on fish collected between dawn and sunset and by estimating the time required for gnathiids to become engorged on host fluids. A model was developed to estimate gnathiid abundance on fish for any given time of day and host size. Fish at dawn had 2.4 times as many gnathiids compared with fish at sunset, indicating that some gnathiids infest fish overnight. Most gnathiids had engorged guts (72–86%); the proportion of empty guts and engorged guts did not differ in three time periods of collection (<0800 h, 0800 to 1100 h, and >1100 h). In the laboratory, gnathiids fed quickly with 75% of gnathiids exposed to fish for 4 h having engorged guts. The short time required for gnathiids to become engorged and the presence of gnathiids with empty guts throughout the day suggests that gnathiids also infest fish during the day. Thus gnathiids eaten by cleaner fish during the day may be replaced by other gnathiids during the day or night suggesting that interactions between gnathiids and cleaner fish are highly dynamic. Accepted: 15 April 1999     

Host specificity of two species of Gnathia (Isopoda) determined by DNA sequencing blood meals

Host specificity data for gnathiid isopods are scarce because the parasitic stages are difficult to identify and host-parasite contact is often brief. We examined two common nocturnal species, Gnathia falcipenis and Gnathia sp. C, collected in light traps from two locations at Lizard Island on the Great Barrier Reef, Australia. Engorged third stage gnathiids were photographed and permitted to moult into adults to allow identification. We compared approximately 580 bp sequences of 16S mtDNA from blood meals with host sequences available on GenBank using BLASTn. Where homology was <98%, familial identity was investigated with neighbour-joining trees. All blood meal sequences (n=60) and homologous fish sequences (n=87) from GenBank were used in a Bayesian analysis, which identified all but three sequences to family. The host frequency distributions used by each species were significantly different; only four host families were shared. No gnathiids fed on elasmobranchs, blennies or apogonids, and most fed on host families whose representatives are typically large. Gnathia sp. C showed a distinct predilection for nemipterids. Gnathia falcipenis often parasitised sand-dwelling families, and unlike sympatric diurnal gnathiid species, it also frequently parasitised pomacentrids. We conclude that G. falcipenis and Gnathia sp. C operate as generalist micropredators with preferences.     

Molecular Characterization of a Parasitic Tapeworm (Ligula) Based on DNA Sequences from Formalin-Fixed Specimens

Museum specimens of Ligula (Pseudophyllidea, Ligulidae), a fish parasite tapeworm, that have been preserved in ethanol or fixed permanently in formalin up to 24 years were used for DNA extraction and molecular characterization. DNA was amplified via PCR from samples collected from different fish hosts that lived in both salt and fresh water bodies in the Chinese Qinghai-Tibet Plateau, Russia, and England. Phylogenetic analyses based on partial nucleotide sequences of the 5'-end of nuclear 28S rRNA gene and the mitochondrial cytochrome c oxidase subunit I (COI) gene support the morphologically based taxonomy that groups the Chinese Ligula within the same species as their Europe counterpart: Lingula intestinalis. No nucleotide variation was detected in either the 28S rRNA gene or the COI gene among the seven plerocercoid samples, suggesting a considerable genetic homogeneity among Ligula from different regions. Our results show that geographic isolation, affinity of hosts, and host habitats are not reliable taxonomic criteria for Ligula classification. Our data also indicate a low genetic diversity in the Ligula prevalent in China and in Europe. Our experiments demonstrate that endogenous DNA from specimens that were subjected to permanent formalin fixation can be routinely amplified from parasitic tapeworms, suggesting that fixation time in formalin may not be a critical factor affecting DNA degradation in such museum specimens.     

Inter- and intraspecific conflicts between parasites over host manipulation

Host manipulation is a common strategy by which parasites alter the behaviour of their host to enhance their own fitness. In nature, hosts are usually infected by multiple parasites. This can result in a conflict over host manipulation. Studies of such a conflict in experimentally infected hosts are rare. The cestode Schistocephalus solidus (S) and the nematode Camallanus lacustris (C) use copepods as their first intermediate host. They need to grow for some time inside this host before they are infective and ready to be trophically transmitted to their subsequent fish host. Accordingly, not yet infective parasites manipulate to suppress predation. Infective ones manipulate to enhance predation. We experimentally infected laboratory-bred copepods in a manner that resulted in copepods harbouring (i) an infective C plus a not yet infective C or S, or (ii) an infective S plus a not yet infective C. An infective C completely sabotaged host manipulation by any not yet infective parasite. An infective S partially reduced host manipulation by a not yet infective C. We hence show experimentally that a parasite can reduce or even sabotage host manipulation exerted by a parasite from a different species.     

Impact of micropredatory gnathiid isopods on young coral reef fishes

The ecological role of parasites in the early life-history stages of coral reef fish, and whether this varies between fish with and without a pelagic phase, was investigated. The susceptibility to, and effect of reef-based micropredatory gnathiid isopods on larval, recently settled, and juvenile fishes was tested using two damselfishes (Pomacentridae): Neopomacentrus azysron, which has pelagic larvae, and Acanthochromis polyacanthus, which does not. When larval and recently settled stages of N. azysron and very young A. polyacanthus juveniles (smaller than larval N. azysron) were exposed to one or three gnathiids, the proportion of infections did not vary significantly among the three host types or between the number of gnathiids to which the fish were exposed. The overall infection was 35%. Mortality, however, differed among the three gnathiid-exposed host types with most deaths occurring in larval N. azysron; no mortalities occurred for recently settled N. azysron exposed to one or three gnathiids, and A. polyacanthus exposed to one gnathiid. Mortality did not differ significantly between larval N. azysron and A. polyacanthus juveniles, failing to provide support for the hypothesis that reef-based A. polyacanthus juveniles are better adapted to gnathiid attack than fish with a pelagic phase. The study suggests that settling on the reef exposes young fish to potentially deadly micropredators. This supports the idea that the pelagic phase may allow young fish to avoid reef-based parasites.     

Phylogeny of nuclear small subunit rRNA genes of hemogregarines amplified with specific primers

Hemogregarines, apicomplexan intracellular blood parasites, are cosmopolitan in distribution and infect a broad range of vertebrate and invertebrate hosts. Molecular phylogenetic studies have been hampered by lack of hemogregarine-specific polymerase chain reaction primers that would allow amplification of parasite, but not host, DNA. A novel method for separating parasite and host 18S rRNA genes has been developed, and new primers that are specific for hemogregarine rRNA genes have been designed. These primers were used to obtain sequences from 4 isolates of hemogregarines of lizards from California, the Caribbean island of Grenada, eastern Australia, and Israel. Combining these results with already published sequences, a preliminary phylogeny of hemogregarines and several other apicomplexan taxa has been created. The hemogregarines form a monophyletic group and appear to be more closely related to coccidia than to Plasmodium species. The difficulty of using 18S genes that have multiple copies in some apicomplexan parasites was explored for systematic studies.     

Comparison of ribosomal DNA sequences of Lamellodiscus spp. (monogenea, diplectanidae) parasitising Pagellus (sparidae, teleostei) in the North Mediterranean Sea: species divergence and coevolutionary interactions

We sequenced DNA fragments from four monogenean species of the genus Lamellodiscus and their three fish host species from the genus Pagellus in the North Mediterranean Sea, in order to estimate the molecular divergence and the coevolutionary interactions in this association. By comparing the ITS1 sequences of the parasites, we assessed their level of interspecific differences and tested the phylogenetic status of Lamellodiscus virgula and Lamellodiscus obeliae, formerly described as two different species. Moreover, we wanted to know if closely related parasites used closely related hosts, to investigate the coevolutionary interactions in this complex. Phylogenetic relationships among Lamellodiscus species were estimated with partial 18S ribosomal DNA sequences while mitochondrial cytochrome-b DNA sequences were used for their fish hosts. The ITS1 sequences appear to be highly variable among Lamellodiscus species, except L.virgula and L.obeliae, suggesting an old divergence time or a rapid molecular evolution within this genus. This fish-parasite association seems to exhibit coevolutionary interactions. L.virgula and L.obeliae are proposed to be a single species on the basis of their almost identical ITS1 sequences.     

Live coral repels a common reef fish ectoparasite

Coral reefs are undergoing rapid changes as living corals give way to dead coral on which other benthic organisms grow. This decline in live coral could influence habitat availability for fish parasites with benthic life stages. Gnathiid isopod larvae live in the substratum and are common blood-feeding parasites of reef fishes. We examined substrate associations and preferences of a common Caribbean gnathiid, Gnathia marleyi. Emergence traps set over predominantly live coral substrata captured significantly fewer gnathiids than traps set over dead coral substrata. In laboratory experiments, gnathiids preferred dead coral and sponge and tended to avoid contact with live coral. When live gnathiids were added to containers with dead or live coral, significantly fewer were recovered from the latter after 24 h. Our data therefore suggest that live coral is not suitable microhabitat for parasitic gnathiid isopods and that a decrease in live coral cover increases available habitat for gnathiids.     

Body size,trophic level,and the use of fish as transmission routes by parasites

Within food webs, trophically transmitted helminth parasites use predator–prey links for their own transfer from intermediate prey hosts, in which they occur as larval or juvenile stages, to predatory definitive hosts, in which they reach maturity. In large taxa that can be used as intermediate and/or definitive hosts, such as fish, a host species’ position within a trophic network should determine whether its parasite fauna consists mostly of adult or larval helminths, since vulnerability to predation determines an animal’s role in predator–prey links. Using a large database on the helminth parasites of 303 fish species, we tested whether the proportion of parasite species in a host that occur as larval or juvenile stages is best explained by their trophic level or by their body size. Independent of fish phylogeny or habitat, only fish body length emerged as a significant predictor of the proportion of parasites in a host that occur as larval stages from our multivariate analyses. On average, the proportion of larval helminth taxa in fish shorter than 20 cm was twice as high as that for fish over 100 cm in length. This is consistent with the prediction that small fishes, being more vulnerable to predation, make better hosts for larval parasites. However, trophic level and body length are strongly correlated among fish species, and they may have separate though confounded effects on the parasite fauna exploiting a given species. Helminths show varying levels of host specificity toward their intermediate host when the latter is the downstream host involved in trophic transmission toward an upstream definitive host. Given this broad physiological compatibility of many helminths with fish hosts, our results indicate that fish body length, as a proxy for vulnerability to predators, is a better predictor of their use by helminth larvae than their trophic level based on diet content.     

Evolutionary divergence in common marine ectoparasites Gnathia spp. (Isopoda: Gnathiidae) on the Great Barrier Reef: phylogeography, morphology, and behaviour

The blood‐feeding juvenile stages of gnathiid isopods are important ectoparasites of marine fishes on the Great Barrier Reef (GBR), and are a major component of the diet of cleaner fishes. We report here that these gnathiids have undergone evolutionary diversification, both geographically and temporally (into diurnally and nocturnally active taxa), which has been accompanied by changes in their morphology and behaviour. To perform this analysis, we sequenced a portion of the nuclear ribosomal ITS2 for 47 gnathiids collected from 29 host fishes of 11 species at three locales spanning 2000 km on the GBR. Maximum parsimony and Bayesian phylogenetic analyses both revealed four major clades. There was some degree of geographical structuring in these clades, but there was no evidence supporting host fish specialization, as gnathiids collected from the skin of different teleost taxa did not resolve into distinct clades. The topology of the phylogeny also implied some structuring that was dependent upon collection time (day or night), so we investigated whether there were also behavioural and morphological differences between taxa active at these different times. Nocturnal gnathiids had significantly longer antennules and larger eyes than diurnal gnathiids – two traits presumably adaptive for nocturnal activity. Behavioural tests showed that both nocturnal and diurnal gnathiids use olfaction and vision while foraging, but that nocturnal gnathiids used olfaction more often in dark conditions, and that they were able to perceive movement under extremely low levels of light. Diurnal gnathiids used vision more effectively when there was some ambient light. Our results thus suggest that both phenotypic and genotypic divergence in gnathiids may be influenced by natural selection acting on ecological traits, such as predator avoidance and host detection. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 94 , 569–587.     

Sex discriminatory effect of the acanthocephalan Acanthocephaloides propinquus on a gobiid fish Gobius bucchichii

Most studies which aim at detecting effects of parasites on fish show that intermediate stages of parasites affect their host while adult parasites are usually less virulent in the final host. We studied the effect of the acanthocephalan Acanthocephaloides propinquus on one of its final hosts, the fish Gobius bucchichii. This study showed that the adult parasites affect host fitness. Moreover, our results showed that at the same level of infection in male and female gobiids, the female's reproductive success was negatively correlated to parasite abundance while the male's reproductive success was not. The negative effects on females includes reductions in gonado-somatic index and egg production. We hypothesized that it might be more difficult for females to compensate the cost of parasitism because of the very high cost of egg production. We discussed these results in terms of host population regulation.     

Interspecific associations among larval helminths in fish

Various processes can generate associations between the larvae of different helminth species in their fish intermediate or paratenic host. We investigated the pairwise associations among larval helminth species in eight different fish populations, using two different coefficients of associations, in order to determine in what situations they are strongest. All helminth species included use the fish studied as either their second intermediate host or their paratenic host, and are acquired by the fish when it ingests an infected first intermediate host. The intensity of infection correlated positively with fish length for most helminth species. Pairs of species which both exhibited positive correlations with fish length tended to be more strongly associated with one another, although this tendency was not pronounced. Similarity in life cycle had a more important influence on pairwise associations. Among the 62 pairwise associations that could be computed, pairs of helminth species that shared both first intermediate hosts and definitive hosts were the most strongly associated, followed by pairs that shared only one other host, and finally by pairs that did not share other hosts. The results suggest that assemblages of larval helminth parasites in fish are not random collections of locally available species, but rather structured packets of larval parasites that travel together along common transmission routes.     

Haustoria in action: Case studies of nitrogen acquisition by woody xylem-tapping hemiparasites from their hosts

Summary This review discusses studies conducted by the author and his colleagues on mistletoes and root hemiparasites native to Western Australia. Morphological characteristics of haustoria are described and their anatomical features are discussed in relation to uptake, transfer, and metabolism of xylem-borne nitrogenous solutes derived from a host. Experimental approaches used include comparisons of xylem sap composition of parasite and host(s), solute pool analyses and enzymatic properties of haustoria, host xylem feeding of¹⁵N-labelled solutes to follow the fate of label in haustoria and body of the parasite, and studies using species-specific nonprotein amino acids to validate successful uptake from hosts or occasional backflow of xylem-borne solutes to a host. Field studies on promiscuous root hemiparasites assess frequencies of exploitation of different hosts.¹⁵N natural abundance assays of host and parasite dry matter demonstrate marked preference bySantalum acuminatum for N₂-fixing as opposed to nonfixing hosts. The ability ofOlax phyllanthi to continue to exploit deep-rooted hosts ranks of importance when xylem water potentials of other hosts go out of sucking range during periods of water stress. Comparisons of xylem sap composition of parasites feeding on different hosts indicate remarkable versatility by haustoria in uptake and utilization of the different major nitrogenous solutes received from these hosts. Solute pools in parasites partly reflect metabolic transformations accentuated by haustoria while also indicating direct throughput from xylem of a host. The review concludes by showing how empirically based modelling techniques can be used to estimate proportional gains of N by parasites from single hosts and repercussions on host growth which accompany such exploitation.Dedicated to Professor Brian E. S. Gunning on the occasion of his 65th birthday     

Monoxenous and heteroxenous parasites of fish manipulate behavior of their hosts in different ways

Adaptive host manipulation hypothesis is usually supported by case studies on trophically transmitted heteroxenous endoparasites. Trematodes and cestodes are among efficient manipulators of fish, their common intermediate hosts. In this review paper, new data on modifications of host fish behavior caused by monoxenous ectoparasitic crustaceans are provided together with a review of effects caused by heteroxenous parasites. Differences in modifications of host behavior caused by heteroxenous and monoxenous parasites are discussed. Manipulation by heteroxenous parasites enhances availability of infected fish to predators--definitive hosts of the parasites. Fine-tuned synchronization of modified anti-predator behavior with a certain phase of the trematode Diplostomum spathaceum development in the eyes of fish, their second intermediate host, was shown. Modifications of behavior are habitat specific. When juvenile salmonids are in the open water, parasites impair their cooperative anti-predator behavior; in territorial bottom-dwelling salmonids, individual defense behavior such as sheltering is the main target of manipulation. It was shown that monoxenous ectoparasitic crustaceans Argulus spp. decreased motor activity, aggressiveness and increased shoal cohesiveness of infected fish. Such a behavior facilitates host and mate searching in these parasites, which often change their hosts, especially during reproduction. Reviewed experimental data suggest that heteroxenous parasites manipulate their host mainly through impaired defense behavior, e.g. impairing shoaling in fish. Alternatively, monoxenous parasites facilitate shoaling that is profitable for both parasites and hosts. Coordination of modified host behavior with the parasite life cycle, both temporal and spatial, is the most convincing criterion of the adaptive value of host manipulation.     

Fluorescent vital labeling to track cestodes in a copepod intermediate host

In experimental studies of host-parasite interactions, it is often important to track parasites in their hosts and to discriminate between individual parasites. We used the fluorescent tracer dyes 7-amino-4-chloromethylcoumarin (CMAC) and, 5-chloromethylfluorescein diacetate (CMFDA) for vital labeling of Schistocephalus solidus (Cestoda) coracidia larvae. Labeling was fast and easy to perform and enabled microscopic detection of parasites appearing as procercoids in the hemocoel of the copepod intermediate host at 3 h after exposure. The label was still visible after 14 days. Extensive controls showed that CMAC (20 microM) labeling did not harm tapeworms or copepods. CMFDA (2 microM) reduced host survival, but the dye concentration can be decreased to avoid this in future studies. The new labeling method presented here has been very useful to track S. solidus parasites. It can be valuable for other parasites also and may be particularly suitable for visualization of individual live macroparasites in invertebrate hosts, for which we are not aware of any other appropriate method.     

Predicting the similarity of parasite communities in freshwater fishes using the phylogeny,ecology and proximity of hosts

Parasite communities tend to be dissimilar in hosts that are geographically, phylogenetically, ecologically and developmentally distant from one another. The decay of community similarity is a powerful and increasingly common method of studying parasite beta diversity, but most studies have examined only a single type of distance. Here, we evaluate distances based on the phylogeny, ecology, spatial proximity and size of hosts, as predictors of the similarity of parasite communities in individual hosts, host populations and host species. We surveyed parasites in six species of fish collected simultaneously from six localities in the St. Lawrence River, Canada, and species in a common group of larval parasites were discriminated using DNA sequences from barcode region of cytochrome c oxidase I. Distances based on the habitat use patterns of host species were good predictors of short‐term, ecological similarity of parasite communities, such as that operating at the scale of the individual host. The genetic distance between host species was associated with almost all types of similarity at all scales, particularly qualitative and phylogenetic similarity of parasite communities at the level of populations and meta‐populations of hosts. The trophic level, diet, spatial proximity and size of hosts were poor predictors of parasite community similarity. The increased taxonomic resolution provided by molecular data increased the explanatory power of regression models, and different factors were implicated when parasite species were distinguished with DNA barcodes than when larval parasites were lumped into morphospecies, as is commonly practiced.     

Effects of parasites on fish behaviour: a review and evolutionary perspective

Fish serve as hosts to a range of parasites that are taxonomically diverse and that exhibit a wide variety of life cycle strategies. Whereas many of these parasites are passed directly between ultimate hosts, others need to navigate through a series of intermediate hosts before reaching a host in (or on) which they can attain sexual maturity. The realisation that parasites need not have evolved to minimise their impact on hosts to be successful, and in many cases may even have a requirement for their hosts to be eaten by specific predators to ensure transmission, has renewed interest in the evolutionary basis of infection-associated host behaviour. Fishes have proved popular models for the experimental examination of such hypotheses, and parasitic infections have been demonstrated to have consequences for almost every aspect of fish behaviour. Despite a scarcity of knowledge regarding the mechanistic basis of such behaviour changes in most cases, and an even lower understanding of their ecological consequences, there can be little doubt that infection-associated behaviour changes have the potential to impact severely on the ecology of infected fishes. Changes in foraging efficiency, time budget, habitat selection, competitive ability, predator-prey relationships, swimming performance and sexual behaviour and mate choice have all been associated with – and in some cases been shown to be a result of – parasite infections, and are reviewed here in some detail. Since the behavioural consequences of infections are exposed to evolutionary selection pressures in the same way as are other phenotypic traits, few behavioural changes will be evolutionarily neutral and host behaviour changes that facilitate transmission should be expected. Despite this expectation, we have found little conclusive evidence for the Parasite Increased Trophic Transmission (PITT) hypothesis in fishes, though recent studies suggest it is likely to be an important mechanism. Additionally, since the fitness consequences of the many behavioural changes described have rarely been quantified, their evolutionary and ecological significance is effectively unknown.Potential hosts may also change their behaviour in the presence of infective parasite stages, if they adopt tactics to reduce exposure risk. Such `behavioural resistance', which may take the form of habitat avoidance, prey selectivity or avoidance of infected individuals, can be viewed as behavioural change associated with the threat of being parasitised, and so is included here. Actually harbouring infections may also stimulate fishes to perform certain types of simple or complex behaviours aimed at removing parasites, such as substrate scraping or the visitation of cleaning stations, although the efficacy of the latter as a parasite removal strategy is currently subject to a good deal of debate.The effects parasites have on shoaling behaviour of host fish have attracted a good deal of attention from researchers, and we have provided a case study to summarise the current state of knowledge. Parasites have been shown to affect most of the antipredator effects of shoaling (such as vigilance, co-ordinated evasion and predator confusion) and can also impair an individual's foraging ability. It therefore seems unsurprising that, in a number of species avoidance of parasitised individuals has evolved which may explain the occurrence of parasite-assorted shoals in the field. Parasitised fish are found more often in peripheral shoal positions and show a reduced tendency for shoaling in some fish species. Given the array of host behaviours that may be changed, the fitness consequences of shoal membership for parasitised hosts and their parasites are not always easy to predict, yet an understanding of these is important before we can make predictions regarding the ecological impact of infections on host fish populations.Clearly, there remain many gaps in our knowledge regarding the effects of parasites on the behaviour of host fish. We believe that a much greater understanding of the importance of infection-associated behaviour changes in fish could be gained from high quality research in comparatively few areas. We have completed our review by highlighting the key research topics that we believe should attract new research in this field.