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.     

Hematozoa of teleosts from Lizard Island, Australia, with some comments on their possible mode of transmission and the description of a new hemogregarine species

Little is known of the blood parasites of coral reef fishes and nothing of how they are transmitted. We examined 497 fishes from 22 families, 47 genera, and 78 species captured at Lizard Island, Australia, between May 1997 and April 2003 for hematozoa and ectoparasites. We also investigated whether gnathiid isopods might serve as potential vectors of fish hemogregarines. Fifty-eight of 124 fishes caught in March 2002 had larval gnathiid isopods, up to 80 per host fish, and these were identified experimentally to be of 2 types, Gnathia sp. A and Gnathia sp. B. Caligid copepods were also recorded but no leeches. Hematozoa, found in 68 teleosts, were broadly hemogregarines of 4 types and an infection resembling Haemohormidium. Mixed infections (hemogregarine with Haemohormidium) were also observed, but no trypanosomes were detected in blood films. The hemogregarines were identified as Haemogregarina balistapi n. sp., Haemogregarina tetraodontis, possibly Haemogregarina bigemina, and an intraleukocytic hemogregarine of uncertain status. Laboratory-reared Gnathia sp. A larvae, fed experimentally on brushtail tangs, the latter heavily infected with the H. bigemina-like hemogregarine, contained hemogregarine gamonts and possibly young oocysts up to 3 days postfeeding, but no firm evidence that gnathiids transmit hemogregarines at Lizard Island was obtained.     

Mancae of the parasitic cymothoid isopod, <Emphasis Type="Italic">Anilocra apogonae</Emphasis>: early life history,host-specificity,and effect on growth and survival of preferred young cardinal fishes

Juvenile parasitic cymothoid isopods (mancae) can injure or kill fishes, yet few studies have investigated their biology. While the definitive host of the adult cymothoids is usually a single host from a particular fish species, mancae may use so-called optional intermediate hosts before settling on the definitive host. Little, however, is known about these early interactions. The cymothoid isopod, Anilocra apogonae, infests the definitive host, Cheilodipterus quinquelineatus. This study examined their host preference among potential optional intermediate hosts. Their effect on the growth and mortality of the young of three apogonid fishes, including the definitive host, was investigated. The number of mancae produced per brood was positively correlated with female length. When given a choice of intermediate hosts, significantly more mancae attached to Apogon trimaculatus (Apogonidae) than to Apogon nigrofasciatus. When presented with Ap. trimaculatus and Pomacentrus amboinensis (Pomacentridae), mancae only attached to Ap. trimaculatus suggesting that mancae may show a taxonomic affiliation with preferred hosts. Mancae fed on all three apogonid species, with C. quinquelineatus being fed on earlier than Ap. trimaculatus and Ap. nigrofasciatus. Mancae feeding frequency, adjusted for fish survival, was lowest on C. quinquelineatus and highest on Ap. trimaculatus. Infested apogonids had reduced growth and increased mortality compared with uninfested fish. A. apogonae mancae can use several species of young apogonid fishes as optional intermediate hosts. Via reduced growth and increased mortality, mancae have the potential to negatively influence definitive host populations and also other young species of apogonid fishes.     

Linking ecology with parasite diversity in Neotropical fishes

A comparative analysis was performed to seek large-scale patterns in the relationships between a set of fish species traits (body size, type of environment, trophic level, schooling behaviour, depth range, mean habitat temperature, geographical range, ability to enter brackish waters and capability of migration) and the diversity of their metazoan parasite assemblages among 651 Neotropical fish species. Two measurements of parasite diversity are used: the species richness and the taxonomic distinctness of a fish's parasite assemblage, including all metazoan parasites, ectoparasites only, or endoparasites only. The results showed that, on this scale, the average taxonomic distinctness of parasite assemblages was clearly more sensitive to the influence of host traits than parasite species richness. Differences in the taxonomic diversification of the parasite assemblages of different fish species were mainly related to the fish's environment (higher values in benthic–demersal species), trophic level (positive correlation with increasing level), temperature (positive correlation with temperature in marine ectoparasites, negative in endoparasites; positive for all groups of parasites in freshwater fishes) and oceanic distribution (higher values in fish species from the Pacific Ocean than those of the Atlantic). The results suggest that, among Neotropical fish species, only certain key host traits have influenced the processes causing the taxonomic diversification of parasite assemblages.     

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.     

Size-related mortality due to gnathiid isopod micropredation correlates with settlement size in coral reef fishes

The transition between the planktonic and the benthic habitat is a critical period for the larvae of many demersal marine organisms. Understanding the potential constraints on the timing of this habitat transition, called settlement, is important to understanding their biology. Size-specific mortality can set the limits on lifestyle and help explain ontogenetic habitat shifts. We examined whether size-based mortality risks after settlement may include micropredation by ectoparasites by testing whether survival of settlement-stage fish varies with fish size when exposed to a reef-associated micropredator. Fish (14 species) were exposed to one blood-sucking gnathiid isopod overnight, with appropriate controls; gnathiid feeding success and survival, and fish mortality were recorded relative to fish size. After adjusting for fish relatedness, we found the relationship between fish mortality and size differed with gnathiid exposure: for gnathiid-exposed fish, the mean mortality of the smallest fish was much higher (57%) than unexposed controls (10%), and decreased to ~0% for fish >12 mm standard length (SL); mortality was almost nil in controls. Thus, a predicted optimal size to switch habitat and reduce mortality risk from micropredation should be >12 mm SL. We then asked what species might be at greater risk and if the steep increase in survival at ~12 mm SL might coincide with settlement at larger sizes among fishes. Across 102 other species (32 families), 61% settled at ≥12 mm SL. After adjusting for relatedness, mean fish settlement size was 15.0 mm and this was not significantly different from 12 mm. Thus, settlement size clusters around the minimum fish size threshold our gnathiid experiment predicted would be large enough to survive a gnathiid encounter. These results suggest micropredators may contribute to size-selective mortality during settlement processes and are consistent with the hypothesis that the pelagic phase provides fish an escape from certain micropredators.     

Vertical distributions of late stage larval fishes in the nearshore waters of the San Blas Archipelago, Caribbean Panama

Light traps were used to describe the vertical distribution of late larval stages of reef fishes in the San Blas Archipelago during three successive new moon periods. Traps were deployed in the lagoon and at an exposed site on the outer reef edge. At each site, two traps were anchored at the surface and two traps just above the bottom. Most families of reef fishes that were abundant in catches displayed clear patterns of depth preference. The larvae of gerrids, pomacentrids and lutjanids were predominantly captured in shallow traps, while gobiids, labrids, apogonids and blenniids were usually collected in deep traps. Studies that used lights to aggregate and collect larval fishes display marked differences in the composition of catches between the Great Barrier Reef (GBR) and the Caribbean. In order to determine whether such results were due to biases inherent in different sampling methods, or to locality-specific patterns of larval behaviour, we simultaneously deployed light traps and dip-netting around lights during three new moons in the San Blas Archipelago. We found that these sampling techniques collected differing components of the larval fish assemblage from the same water mass. However, there remains good evidence for the existence of locality-specific responses to light in older larval stages, suggesting that broad generalisations about patterns and causes of vertical distributions may be difficult to achieve.     

Blood feeding in juvenile Paragnathia formica (Isopoda: Gnathiidae): biochemical characterization of trypsin inhibitors, detection of anticoagulants, and molecular identification of fish hosts

The 3 post-marsupial juvenile stages of the gnathiid isopod, Paragnathia formica, are haematophagous ectoparasites of fishes that may, in heavy infestations, cause host mortality. Protein digestion in fed stage 3 juveniles is accomplished by cysteine proteinases, but what bioactive compounds attenuate host haemostatic, inflammatory and immunological responses during feeding is unknown. Trypsin inhibitory activity and anticoagulant activity were detected in crude extracts of unfed P. formica stage 1 juveniles; fractionation of stage 1 crude extracts by ion exchange chromatography resulted in 3 preparations each displaying these bioactivities. Further characterization revealed anti-thrombin activity in 2 of these preparations, whilst the third displayed the strongest anticoagulant activity that targeted a factor of the intrinsic coagulation pathway. Three trypsin inhibitors (18 kDa, 21 kDa, and 22 kDa) were also detected using reverse zymography. In parallel, homogenates of fed stage 2 and 3 juveniles were used to identify their fish hosts by amplifying the 16S mitochondrial rDNA and 18S genomic rDNA vertebrate gene regions. Blood from at least 4 fish families had been ingested by separate individuals during feeding. This study demonstrates that trypsin inhibitors and anticoagulants are present in P. formica juveniles which could suppress host haemostatic, inflammatory and immunological responses during feeding, and that juveniles are not host specific.     

Group-living and the richness of the parasite fauna in Canadian freshwater fishes

Summary An increased transmission of ectoparasites among individual animals is considered to be an inevitable cost of living in groups, since several kinds of ectoparasites require close proximity between large numbers of hosts for successful transmission. However, we do not know whether individuals belonging to group-living species incur a greater risk of ectoparasitism than individuals of solitary species. Here, using published data from 3 families (60 species) of Canadian freshwater fishes, I test the hypothesis that group-living species are host to more species of contagious ectoparasites (copepods and monogeneans) than solitary host species. As the different fish species have been studied with varying intensity, I used the mean number of parasite species recorded per study as a standard measure of parasite numbers. Multiple regression analyses were performed separately for each family to determine the effects of group-living and of 3 other variables (host size, age, and range) on the richness of the recorded parasite fauna. Once the effects of the other variables were removed, I found no significant effect of sociality on the richness of the parasite fauna per fish species, for contagious ectoparasites and other types of parasites. Neither of the other variables had any influence on the numbers of parasite species per fish species. These results suggest that a richer ectoparasite fauna is not a cost of group-living in fishes.     

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.     

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.     

Diel infestation dynamics of gnathiid isopod larvae parasitic on Caribbean reef fish

Infestation dynamics of parasitic gnathiid isopods on Caribbean reefs were studied throughout the 24-h diel cycle. Gnathiid infestation on caged longfin damselfish (Stegastes diencaeus) peaked strongly at dawn, remained low during the remainder of the day, and increased again at night until about midnight. Gnathiids were less abundant during the pre-dawn period. Peak loads on fish retrieved at dawn were the highest reported in any study thus far. The dawn peak consisted almost exclusively of individuals from the smallest size class, whereas nocturnal activity consisted almost exclusively of individuals of the largest size class. Because of the high rates of infestation at night and dawn, and the high variation in parasite loads on fish collected during that time, reduction of parasite infestation may play an important role in the selection of nocturnal and crepuscular shelter holes and settlement sites by reef fishes.     

A simple molecular technique for identifying marine host fish by sequencing blood-feeding parasites

Gnathiid isopods are common ectoparasites of fish on the Great Barrier Reef, Australia. While screening for appropriate markers for phylogenetic studies of gnathiids, we found that primers for 12S and 16S rDNA preferentially amplified the host fish DNA instead of gnathiid DNA. This amplification occurred even when using gnathiids that were not engorged with host blood and adult gnathiids that do not feed on fish blood. This method could be used in host-parasite studies to identify hosts without having to sample parasites directly from the host (which can be costly and requires considerable skill in a marine environment). Target ribosomal DNA sequences can be amplified from total DNA extracted from parasites that are captured in funnel traps or plankton tows. Sequence data from these can be used to identify the hosts that gnathiids were feeding on before capture.     

Phylogenetic relationships and evolutionary history of the reef fish family Labridae

The family Labridae (including scarines and odacines) contains 82 genera and about 600 species of fishes that inhabit coastal and continental shelf waters in tropical and temperate oceans throughout the world. The Labridae (the wrasses) is the fifth largest fish family and second largest marine fish family, and is one of the most morphologically and ecologically diversified families of fishes in size, shape, and color. Labrid phylogeny is a long-standing problem in ichthyology that is part of the larger question of relationships within the suborder Labroidei. A phylogenetic analysis of labrids was conducted to investigate relationships among the six classical tribes of wrasses, the affinities of the wrasses to the parrotfishes (scarines), and the broad phylogenetic structure among labrid genera. Four gene fragments were sequenced from 98 fish species, including 84 labrid fishes and 14 outgroup taxa. Taxa were chosen from all major labrid clades and most major global ocean regions where labrid fishes exist, as well as cichlid, pomacentrid, and embiotocid outgroups. From the mitochondrial genome we sequenced portions of 12S rRNA (1000 bp) and 16S rRNA (585 bp), which were aligned by using a secondary structure model. From the nuclear genome, we sequenced part of the protein-coding genes RAG2 (846 bp) and Tmo4C4 (541 bp). Maximum likelihood, maximum parsimony, and Bayesian analyses on the resulting 2972 bp of DNA sequence produced similar topologies that confirm the monophyly of a family Labridae that includes the parrotfishes and butterfishes and strong support for many previously identified taxonomic subgroups. The tribe Hypsigenyini (hogfishes, tuskfishes) is the sister group to the remaining labrids and includes odacines and the chisel-tooth wrasse Pseudodax moluccanus, a species previously considered close to scarines. Cheilines and scarines are sister-groups, closely related to the temperate Labrini, and pseudocheilines and cheilines are split in all phylogenies. The razorfishes (novaculines) and temperate pseudolabrines form successive sister clades to the large crown group radiation of the Julidini. The cleaner wrasses (Labrichthyini) are nested within this radiation and several julidine genera do not appear to be monophyletic (e.g., Coris and Halichoeres). Invasion of temperate water by this predominantly tropical group has occurred multiple times and the reconstruction of biogeography assuming an Indo-Pacific ancestor results in five different lineages invading the Atlantic/Caribbean region. Functional novelties in the feeding apparatus have allowed labrid fishes to occupy nearly every feeding guild in reef environments, and trophic variation is a central axis of diversification in this family.     

Effects of Habitat Connectivity on the Abundance and Species Richness of Coral Reef Fishes: Comparison of an Experimental Habitat Established at a Rocky Reef Flat and at a Sandy Sea Bottom

Coral reef fish assemblages are widely recognized for the coexistence of numerous species, which are likely governed by both coral diversity and substratum complexity. However, since coral reefs provide diverse habitats due to their physical structure and different spatial arrangements of coral, findings obtained from an isolated habitat cannot necessarily be applied to fish assemblages in other habitats (e.g. continuous habitats). The aim of this study, therefore, was to determine by a field experiment whether habitat connectivity (spatial arrangement of coral colonies) affects abundance and species richness of fishes in an Okinawan coral reef. The experiment consisted of transplanted branching coral colonies at a 4m×8m quadrat at both a rocky reef flat and sandy sea bottom. Generally, the abundance of fishes was greater at the sandy sea bottom, especially for three species of pomacentrids, one species of labrids, one species of chaetodontids and two species of apogonids. Species–area curves showed that the species richness of fishes was significantly greater in the quadrat at the sandy sea bottom at 3, 6 and 9 months after the start of the experiment. The rate of increase in abundance of fishes per area was significantly greater in the quadrat at the sandy sea bottom over the study period. The results of rarefaction analyses showed that the rate of increase in species richness per abundance was significantly higher in the quadrat at the sandy sea bottom in the juvenile settlement period, indicating that the magnitude of dominance by particular species was greater at the sandy sea bottom habitat. Our findings suggest that habitat connectivity affects the abundance and species richness of coral reef fishes, i.e. the isolated habitat was significantly more attractive for fishes than was the continuous habitat. Our findings also suggest that the main ecological factors responsible for organization of fish assemblage at a continuous habitat and at an isolated habitat are different.     

Structure of the parasite communities of a coral reef fish assemblage (Labridae): testing ecological and phylogenetic host factors

The role of ecological and phylogenetic processes is fundamental to understanding how parasite communities are structured. However, for coral reef fishes, such information is almost nonexistent. In this study, we analyzed the structure of the parasite communities based on composition, richness, abundance, and biovolume of ecto- and endoparasites of 14 wrasse species (Labridae) from Lizard Island, Great Barrier Reef, Australia. We determine whether the structure of the parasite communities from these fishes was related to ecological characteristics (body size, abundance, swimming ability, and diet) and/or the phylogenetic relatedness of the hosts. We examined 264 fishes from which almost 37,000 individual parasites and 98 parasite categories (types and species) were recorded. Gnathiid and cestode larvae were the most prevalent and abundant parasites in most fishes. Mean richness, abundance, and biovolume of ectoparasites per fish species were positively correlated with host body size only after controlling for the host phylogeny, whereas no such correlation was found for endoparasites with any host variable. Because most ectoparasites have direct transmission, one possible explanation for this pattern is that increased space (host body size) may increase the colonization and recruitment of ectoparasites. However, endoparasites generally have indirect transmission that can be affected by many other variables, such as number of prey infected and rate of parasite transmission.     

PREFERENCE HIERARCHIES AND THE ORIGIN OF GEOGRAPHIC SPECIALIZATION IN HOST USE IN SWALLOWTAIL BUTTERFLIES

Four allopatric populations of the widely distributed western anise swallowtail butterfly, Papilio zelicaon, use different plant genera as hosts, but simultaneous choice experiments showed that these populations have diverged only slightly in oviposition preference. Of the four populations—two from southeastern Washington State, one from coastal southwestern Washington State, and one from central California—three use hosts that are not available to any of the others. Although variation for the degree of preference for particular plant species occurred within and among populations, all four populations ranked hosts in the same overall order. Monophagy on a local, low-ranking host outside the range of high-ranking hosts did not necessarily lead to the loss of preference for those high-ranking hosts, thereby indicating that the high-ranking hosts would still be accepted, and in some cases even preferred, if a population encountered them again. Hence, the overall preference hierarchy among P. zelicaon populations appears to be evolutionarily conservative. Analyses of differences among families within the California population indicated that increased preference for some hosts is inversely correlated, whereas preference for other hosts may be uncorrelated. Positive correlations may also occur but were not observed among the plant species tested. Overall, the results indicate local monophagy on different plant species in P. zelicaon has not involved major reorganizations in the preference hierarchy of ovipositing females, even in populations that may have fed on a low-ranking host for many generations. Instead, small increases in preference for local hosts have occurred within an evolutionarily conservative preference hierarchy.     

Local phylogenetic divergence and global evolutionary convergence of skull function in reef fishes of the family Labridae

The Labridae is one of the most structurally and functionally diversified fish families on coral and rocky reefs around the world, providing a compelling system for examination of evolutionary patterns of functional change. Labrid fishes have evolved a diverse array of skull forms for feeding on prey ranging from molluscs, crustaceans, plankton, detritus, algae, coral and other fishes. The species richness and diversity of feeding ecology in the Labridae make this group a marine analogue to the cichlid fishes. Despite the importance of labrids to coastal reef ecology, we lack evolutionary analysis of feeding biomechanics among labrids. Here, we combine a molecular phylogeny of the Labridae with the biomechanics of skull function to reveal a broad pattern of repeated convergence in labrid feeding systems. Mechanically fast jaw systems have evolved independently at least 14 times from ancestors with forceful jaws. A repeated phylogenetic pattern of functional divergence in local regions of the labrid tree produces an emergent family-wide pattern of global convergence in jaw function. Divergence of close relatives, convergence among higher clades and several unusual 'breakthroughs' in skull function characterize the evolution of functional complexity in one of the most diverse groups of reef fishes.     

Evaluation of gill nets, fyke nets, and mark-recapture methods to estimate the number of Hirudinea and Crustacea on fish.

Twenty species of fishes (n = 20,759) were collected from Dauphin Lake, Manitoba, Canada, to determine the types and numbers of ectoparasites they harbored. Counts of ectoparasites on fishes collected with different gear were compared to evaluate different methods of collection and to estimate rates of recruitment of ectoparasites by fishes. Ectoparasites were found on 11 species of fishes and the majority of these were parasitic leeches (Myzobdella moorei, Cystobranchus verilli, and Placobdella montifera) and parasitic Crustacea (Argulus appendiculosus and Lernaea cyprinacea). Some fishes also were infested by neascus-type metacercariae (blackspot) or had tumors (lymphocystis). The prevalence of ectoparasites was correlated with the abundance, feeding habits, and spatial distribution of fish species. Argulus appendiculosus and blackspot were more prevalent on benthic fishes, whereas M. moorei and tumors were more prevalent on limnetic fishes. Mark-recapture records showed that fishes occupying shallow (less than or equal to 1.5 m) water had a higher prevalence of infestation and 28 of 29 infected fishes caught by gill nets were captured in shallow water. Placobdella montifera was the only ectoparasite found on fishes from deep (1.5-3.5 m) water and the only species that was acquired by fishes previously released with no ectoparasite (2 of 239 fishes). The littoral zone (less than or equal to 1.5 m) comprises only 14% of the surface area and 3% of the volume of Dauphin Lake, yet 72% of all gill-netted fishes harboring ectoparasites were collected there. Intensities of ectoparasites estimated from gill net and pound net samples were similar, but prevalence of ectoparasites estimated from samples obtained with gill nets was lower.     

Feeding preferences and performance of an aquatic lepidopteran on macrophytes: plant hosts as food and habitat

Although host preferences in phytophagous insects may be generated by several factors, few studies have simultaneously examined several potential host choice determinants. In this study we tested the impact of the following potential host choice determinants on host preference of the semi-aquatic lepidopteran Munroessa gyralis (Pyralidae): growth on different host plants; protein content, polyphenolic content, toughness, and chemical extracts of different host plants; prior feeding experience; and predation pressure on the caterpillar by fishes. Two water lilies, Brasenia schreberi and Nymphaea odorata, were preferred in cafeteria-style feeding experiments over 14 other species of vascular plants. The most preferred water lily (Brasenia) also afforded the fastest growth relative to three other species on which growth was measured. Feeding preferences across species were unrelated to protein content, polyphenolic content, or toughness. Domiciles constructed by caterpillars from leaf fragments were protective from field assemblages of fishes, but domiciles made from preferred or unpreferred host species conferred no significant protection from fish in the laboratory. Caterpillars responded positively to chemical cues of water lilies, and prior feeding experience increased preference for an otherwise unpreferred water lily (Nuphar advena) within the life-span of individual caterpillars. M. gyralis is a generalist herbivore exhibiting modest preference induction and preferences for and among members of the family Nymphaeaceae. Our results suggest that relative growth rates, chemical cues, and previous feeding experience are important factors determining feeding preference. Protein content, polyphenolic content, and toughness appear less important, and the importance of fish predators remains in question. As pupation seems to occur exclusively on Nymphaea, we suggest that host use may be restricted due to life-stage-specific developmental constraints that are not apparent from the results of growth or preference assays. It is currently unknown how often specific life-stages may restrict host use, but our work suggests this as a potentially important area of inquiry.