Relationship between intermediate host taxon and infection by nematodes of the genus Rhabdochona

We describe the intermediate and definitive hosts of the fish nematodes Rhabdochona coronacauda and R. denudata honshuensis and discuss the relationships between parasitism and the feeding habitats of their intermediate hosts. We found that the principal intermediate hosts of the two nematodes were filter-feeding mayflies of the genera Ephemera, Photamanthus and Isonychia. Ephemera strigata seemed to be the most important intermediate host of these nematodes. Adult R. coronacauda were found mainly in Hemibarbus longirostris and Rhinogobius flumineus, which are benthic fishes that feed on benthic aquatic insects, including E. strigata. For R. coronacauda, therefore, the feeding habits of the definitive hosts facilitate host alternation by this species. However, adult R. denudata honshuensis were found in cyprinids. In particular, Zacco temmincki was the principal natural definitive host in our study area. Since Z. temmincki is a swimming predator, E. strigata nymphs that burrow in the substrate are not the main prey of this species. This indicates that the transmission of R. denudata honshuensis hardly occurs from E. strigata nymph to Z. temmincki, suggesting another, unknown transmission route.     

Observations on the biology of Rhabdochona kidderi texensis, a parasite of North American cichlids

An examination of a sample of benthic invertebrates collected from the Upper San Marcos River in southwestern Texas, USA in September 1999 revealed that the nymph of the ephemeropteran Tricorythodes curvatus served as natural intermediate host of the nematode Rhabdochona kidderi texensis (Nematoda: Rhabdochonidae), an intestinal parasite mainly of the Rio Grande perch (Cichlasoma cyanoguttatum) in this locality; the prevalence of the parasite's third- and fourth-stage larvae in mayflies was 6.8% with the intensity of 1-2 larvae per nymph. Live R. kidderi texensis eggs collected from nematodes recovered from C. cyanoguttatum in Texas were transported to the Czech Republic, where they were used to experimentally infect nymphs of the palaearctic mayfly species Paraleptophlebia submarginata; the development of infective third- and fourth-stage larvae in this experimental intermediate host was completed after approximately 10 days at 19 degrees C. Infected nymphs were fed to aquarium-reared fishes, four Cichlasoma nigrofasciatum and one Oreochromis niloticus, of which only three of the former became infected. The last (fourth) moult of a male nematode was observed in C. nigrofasciatum 23 days p.i. and adult males and gravid females with not fully mature (non-embryonated) eggs in uteri on days 40 and 51 p.i. The prepatent period of R. kidderi texensis is approximately two months.     

Observations on the occurrence and maturation of the nematode Rhabdochona denudata (Dujardin, 1845) in Chub, Leuciscus cephalus (L.), of the Rokytná River, Czechoslovakia

From October 1985 until July 1987, the seasonal dynamics of Rhabdochona denudata in its principal definitive host, the chub (Leuciscus cephalus), was studied in the Rokytná River (the Danube basin), Czechoslovakia. Prevalence (overall prevalence 77%) and intensity of infection (1-59 nematodes per fish) in the fish were high all year, with both abiotic and biotic factors influencing the fluctuations. Maximum values of prevalence (82-87%) were found in the fish with body length exceeding 20 cm; the mean intensity was highest in the size-group of fish 20-25 cm long, whereas its values gradually decreased in larger fish, this being due to the choice of food by fish of different size-groups. In contrast to the situation in other localities, an annual maturation cycle was not evident and egg-producing female nematodes were present in fish throughout the year. New infections in fish were most common in summer but occurred all year round. This life history pattern seems to be mainly due to the availability of infected ephemeropteran intermediate hosts during all seasons, in this case the nymphs of Ephemera danica, a species noted for its two-year development. In this locality, specifically not determined Rhabdochona larvae were recorded from the mayfly nymphs Ephemera danica (0.4%), Ecdyonurus aurantiacus (7%), and Caenis macrura (10%). The diet of the chub included mayfly nymphs at all seasons.     

Occurrence and maturation of Rhabdochona gnedini (Nematoda: Rhabdochonidae) in the barbels of the Sousa River, Portugal

Data on the seasonal occurrence and maturation of Rhabdochona gnedini Skrjabin, 1946 in its definitive host, the barbel Barbus bocagei, from the Sousa River, in northern Portugal, are presented. Monthly samples were taken from November, 1998 to October, 1999. A total of 178 fish specimens were inspected. R. gnedini occurred with high prevalences during whole year, attaining the highest values of intensity in summer (overall prevalence and mean intensity 85% and 31, respectively). A clear annual cycle of maturation was observed; females with mature eggs occurred in spring and summer. Unidentified Rhabdochona larvae were recorded from the trichopteran larvae Hydropsyche sp. (prevalence and intensity 29% and one to 34, respectively). These larvae are presumptive intermediate host for R. gnedini.     

New species of Rhabdochona Railliet, 1916 (Nemata: Rhabdochonidae) from Rainbow Trout in California Streams

Three new species of Rhabdochona Railliet, 1916 are described and illustrated from Salmo gairdneri Richardson (rainbow trout) in freshwater streams in California: Rhabdochona californiemis n. sp., R. paxmani n. sp., and R. satmonis n. sp. Rhabdochona californiensis n. sp. is characterized by 14 anteriorly directed teeth in the prostome, egg devoid of filaments or floats, male and female tail terminus with a single mucro, left (long) spicule slender with a moderate distended podoid terminal end, spicular ratio 1:3.8. Rhabdochona paxmani n. sp. is characterized by 10 teeth in the prostome, eggs with polar floats, left (long) spicule slender with podoid terminus distended and having a minute subterminal spine; right spicule with prominent gorgeret (barb), spicular ratio 1:4.3, male and female tail terminus with a cuticular conical rounded short projection. Rhabdochona salmoni, n. sp. is characterized by 10 teeth anteriorly directed in the prostome, eggs with polar floats, left spicule slender with a distended podoid terminus; right spicule with a sharply indented gorgeret, spicular ratio 1:4.3, male and female tail terminus with a conical or rounded tip.     

Observations on two Rhabdochona species (Nematoda: Rhabdochonidae) from freshwater fishes in Argentina, including description of Rhabdochona fabianae n. sp

Helminthological examination of specimens of the freshwater fishes Bryconamericus iheringi Boulenger, 1887 (Characidae) and Jenynsia multidentata (Jenyns, 1842) (Anablepidae), collected from Medina River, Province of Tucuman, Argentina, revealed the presence of 2 species of parasitic nematodes, Rhabdochona fabianae n. sp. and Rhabdochona acuminata (Molin, 1860) (Rhabdochonidae). Rhabdochona fabianae n. sp. was characterized by the presence of 16 anterior teeth in the prostom and filamented eggs in females. Rhabdochona acuminata was recorded for the first time in the north of Argentina.     

Consequences of tadpole shrimp predation on mayflies in some Californian ponds

SUMMARY. 1. During two, 1-month studies, the abundance of mayfly nymphs, Callibaetis californicus Banks, in small ponds that contained the tadpole shrimp, Triops longicaudatus (LeConte), often was significantly lower than in ponds without tadpole shrimps. Mayfly abundance was either positively correlated or not significantly associated with that of predacious insects.
2. Larger mayfly nymphs were a greater fraction of the population in ponds with Triops than in ponds without tadpole shrimps. Between 2 and 3 weeks after ponds were flooded, the smallest mayfly nymphs were absent from ponds in which T. longicaudatus densities were >5 individuals per square metre during the summer and >15 individuals per square metre during the autumn.
3. In prey-choice trials, T. longicaudatus consumed predominantly small C. californicus nymphs (mean headwidth ± 1 SD: 0.44 ± 0.05 mm). After encounters with tadpole shrimps, small mayfly nymphs moved relatively shorl distances, were easily overtaken, and were, consequently, more susceptible to tadpole shrimps than larger-sized nymphs.     

Rhabdochona parastromatei sp.n. (Nematoda, Rhabdochonidae) from the Fish Parastromateus niger (Bleeker) of the Karachi Coast

Rhabdochona (Rhabdochona) parastromatei sp.n. is characterized by its relatively large body size, 16 teeth in pros tome, right spicule 0.48-0.50 mm and left 0.10-0.12 mm in length, 9–10 pairs of caudal papillae, a single adanal papilla, eggs in female 0.029-0.044 by 0.018-0.025 mm in size, and by having a marine fish host.     

Changing Abundance of Hexagenia Mayfly Nymphs in Western Lake Erie of the Laurentian Great Lakes: Impediments to Assessment of Lake Recovery?

After an absence of 40 years, mayfly nymphs of the genus Hexagenia were found in sediments of western Lake Erie of the Laurentian Great Lakes in 1993 and, by 1997, were abundant enough to meet a mayfly‐density management goal (ca. 350 nymphs m^—2) based on pollution‐abatement programs. We sampled nymphs in western Lake Erie and Lake St. Clair, located upstream of western Lake Erie, to determine the importance of seasonal abundance and life‐history characteristics of nymphs (e.g., emergence and recruitment) on density estimates relative to the mayfly‐density management goal. Two types of density patterns were observed: (1) densities were relatively high in spring and gradually decreased through late summer (observed in Lake Erie and Lake St. Clair in 1997 and Lake St. Clair in 1999) and (2) densities were relatively high in spring, gradually decreased to mid summer, abruptly decreased in mid summer, and then increased between summer and late fall (Lake Erie and Lake St. Clair in 1998 and Lake Erie in 1999). Length‐frequency distributions of nymphs and observations of adults indicate that the primary cause for the two density patterns was attributed to failed (first pattern) and successful (second pattern) reproduction and emergence of nymphs into adults in mid summer. Gradual declines in densities were attributed to mortality of nymphs. Our results indicate that caution should be used when evaluating progress of pollution‐abatement programs based on mayfly densities because recruitment success is variable both between and within years. Additionally, the interpretation of progress toward management goals, relative to the restoration of Hexagenia populations in the Great Lakes and possibly other water bodies throughout the world, is influenced by the number of years in which consequtive collections are made.     

Growth responses of littoral mayflies to the phosphorus content of their food

We examined how mayfly growth rates and body stoichiometry respond to changing phosphorus (P) content in food. In two experiments, mayfly nymphs were given high or low quantities of food at different carbon:phosphorus (C:P) ratios and their growth was measured. Low food quantity resulted in negative growth rates in both experiments, regardless of food P content. However, under high food availability, mayfly growth was affected by the type of food eaten, with low C:P ratio food producing more rapid growth. In addition, mayfly growth increased somewhat when P-poor food was artificially enriched with inorganic P although this effect was not statistically significant. Mayfly body P content was inversely related to body size but increased in animals fed artificially P-enriched food. A model was constructed to simulate mass balance constraints on mayfly growth imposed by the relative supply of two elements (C and P) in food. The model shows that mayfly growth should be limited by food P content at moderately low C:P ratios ( c . 120, by mass). Given high C:P ratios (mean c.  270, by mass) in periphyton from oligotrophic boreal lakes, our experimental and theoretical results indicate that stoichiometric constraints are important factors affecting benthic food webs in lakes from the Canadian Shield and perhaps in other systems with similarly high C:P ratios in periphyton.     

Predator detection and avoidance by lotic mayfly nymphs of different size

We studied antipredatory responses of lotic mayfly (Baetis) nymphs in a factorial experiment with four levels of fish presence: (1) a freely foraging fish (the European minnow,Phoxinus phoxinus), (2) a constrained fish, (3) water from a fish stream, (4) water from a fishless stream. LargeBaetis nymphs drifted mainly during night-time in treatments involving either the chemical or actual presence of fish, whereas no diel periodicity was observed when the water was not conditioned with fish odour. The response was strongest when the fish was uncaged, which suggests that visual or hydrodynamic cues are needed in addition to chemical ones for an accurate assessment of predation risk. Fish presence had no effect on the drift rates of small nymphs. Instead, they increased their refuge use in the presence of a live fish. Chemical cues alone did not have any effect on the refuge use of any of theBaetis size classes. Our results indicate active drift entry by mayfly nymphs. Because predation pressure is spatially and temporally variable, nymphs must sample the environment in order to locate predator-free areas or areas with low predation risk. Drifting should be the most energy-saving way to do this. To avoid the risk from visually feeding fish, large individuals can sample safely (i.e. enter drift) only at night-time, while the small ones can also do this safely during the day. We suggest that, contrary to some earlier assumptions, mayfly drift is not a fixed prey response. Instead,Baetis nymphs are able to assess the prevailing predation pressure, and they adjust their foraging behaviour accordingly.     

Rhabdochona guerreroensis n. sp. (Nematoda: Rhabdochonidae) from the intestine of the gobiid Sicydium multipunctatum from fresh waters in Mexico

Rhabdochona guerreroensis n. sp. is described on the basis of specimens recovered from the intestine of Sicydium multipunctatum (Regan) (Gobiidae, Perciformes) from 2 localities in Mexico. This species is characterized by 12 teeth in the prostom (occasionally 11), asymmetrical numbers of postanal papillae, a broad distal tip of the larger (left) spicule with a small bifurcation covered by narrow cuticular membrane, a smaller (right) spicule with a dorsal barb on its distal tip, and filamented eggs.     

The effects of prey profitability and habitat complexity on the foraging success and growth of stonefly (Plecoptera) nymphs

• 1 The feeding behaviour of nymphs of the perlid stonefly Dinocras cephalotes when presented with two different mayfly prey types on different substrates was examined in the laboratory.
• 2 Most of the evidence suggested that Dinocras cephalotes did not choose between Baetis rhodani and Rhithrogena semicolomta as, under identical circumstances: (i) the survival of both prey types was the same; (ii) the predator's overall searching behaviour was similar in the presence of either mayfly, although B. rhodani was pursued more, especially on complex substrates; (iii) the number of attacks on both prey types was the same; and (iv) the intake of dry weight prey biomass over 24 h remained the same, irrespective of prey species.
• 3 The predation efficiency (no. of prey captured/no, of encounters) of D. cephalotes decreased with increasing substrate complexity—but only with R. semicolorata as prey, and also when more than one stonefly was present—but only when B. rhodani was the prey. Predator efficiency was greater at low densities of R. semicolorata but highest at a medium density of B. rhodani.
• 4 Increase in the body weight of D. cephalotes was greater when nymphs were fed a monospecific diet of R. semicolorata compared with one of B. rhodani. This was despite the findings that: (i) nymphs of B. rhodani contained 31.3% more protein (per mg dry body weight) than nymphs of R. semicolorata, whereas the latter contained 81.0% more chitin; and (ii) nymphs of R. semicolorata took 2.7 times as long to consume and almost twice as long to pass through the predator's gut. The longer gut passage time may have allowed a greater assimilation efficiency which may have led to the observed better growth.
• 5 Dinocras cephalotes is thought to feed opportunistically, with a certain daily food biomass intake as its primary need rather than any specific prey preference. Foraging decisions, or opportunities, affect the fitness of individual nymphs and this may be reflected in the wide size range of contemporary nymphs found in field populations. Variation in foraging efficiency by predators and evasion success by prey, across substrate types, is thought to contribute to the well-known microdistribution patterns of species observed in lotic communities.

     

Mayfly fauna (Ephemeroptera) and the biology of the species Potamanthus luteus (L.) in a warmed stretch of the Oslava River

Cooling waters of the Oslavany power station show a depressive effect on the species diversity of mayfly nymphs. Only seven of 35 taxons found in the study area occurred in all localities. In the warmed stretch of the river, the abundance and biomass of mayfly nymphs as well as the production of the most important speciesPotamanthus luteus decreased; a gradual regeneration of the disturbed mayfly population could be observed downstream. In Locality 5, situated 3.5 km downstream from the mouth of cooling waters, the species diversity, abundance, and biomass of mayflies were 69, 30, and 66 per cent, resp., of values found in Locality 1 with natural temperatures; the production ofPotamanthus luteus was 70 per cent of that found in Locality 1.     

The life history of Plagioporus shawi (McIntosh 1939) (Trematoda:Opecoelida), an intestinal parasite of salmonid fishes.

Podocotyle shawi is transferred to the genus Plagioporus. The miracidium develops and hatches in 11 to 14 days at 22 to 25 C. The operculate snail, Lithoglyphus virens, serves as first intermediate host. Mother and daughter sporocysts develop, the latter producing cotylomicrocercous cercariae within 90 to 130 days. Cercariae encyst in the amphipod, Hyalella azteca; stonefly nymphs of the genus Arcynopteryx; mayfly nymphs of the genera Heptagenia and Paraleptophlebia; chironomid larvae of the genera Chironomus, Polypedilum, Phaenopsectra, Ablabesmyia, and Psectrocladius and caddis larvae of the genera Brachycentrus, Limnephilus, and Hydropsyche. Metacercariae 60 to 100 days old were fed to young steelhead trout, Salmo gairdneri, and partially developed specimens of P. shawi were recovered 26 days later. Stonefly nymphs are believed to be the most important second intermediate host in the natural environment. The parasite seems to be restricted to the Pacific Northwest of the United States and has been reported from the silver salmon, Oncorhynchus kisutch; kokanee, O. nerka; cutthroat trout, Salmo clarki; steelhead trout, S. gairdneri; and the mountain whitefish, Prosopium williamsoni.     

Variation in the sex ratio of Rhabdochona fortunatowi (Spirurida: Rhabdochonidae) in Capoeta capoeta gracilis (Cypriniformes: Cyprinidae), relative to levels of infection, host size and temperature

In the present study, 81 specimens of Capoeta capoeta gracilis (Keyserling, 1861) were collected from the Shiroud River, the Caspian drainage basin, Iran in June and October 2007. The fish were examined for infection with the nematode Rhabdochona fortunatowi (Dinnik, 1933). Nematodes were counted and sexed, then the sex ratio and intestinal site preference of the R. fortunatowi individuals and the intersexual interactions of the parasites were investigated. Significant differences were observed in the prevalence and intensity of infection in relation to host size and sampling season, but not host sex. Parasite burden (mean intensity) was higher in October than in June and the results revealed a decrease in female-to-male sex ratio (FMR) in the fish in October compared to the ones in June. In other words, as the mean intensity of infection of R. fortunatowi increases, the proportion of male worms increases, too. The female-biased sex ratio in June was replaced by a male-biased one in October. Nematode distribution rose to a significant peak in the mid-region of the fish intestine relative to the anterior and posterior parts. Some changes in the distribution of male and female worms in different parts of the host intestine were also observed during June and October. Density-dependent selection and intersexual competition seem to be the main factors driving such a shift in the sex ratio and its variation in different parts of the host intestine.     

The infection of nymphal Baetisbicaudatus by the mermithid nematode Gasteromermis sp.

1. This study reports the infection in nymphs of a bivoltine mayfly host ( Baetis bicaudatus ) in a high-elevation watershed by the mermithid nematode Gasteromermis sp. Infection by Gasteromermis causes mortality in two ways. Fifty per cent of the infections do not successfully develop beyond the initial stage of penetration and result in the early death of both host and parasite.
2. Infected hosts that survive this initial stage are rendered completely sterile by the infection (reproductively dead). In addition to complete sterility, the emergence size of parasitized nymphs is reduced and development time lengthened compared with unparasitized nymphs.
3. Parasite infection levels are stable from year to year at one site, but with a higher incidence of infection in the mayfly summer generation. Size differences between the generations at the time of infection may account for their different susceptibilities.
4. Within a year infection levels vary seasonally and spatially from 1 to 71%. Seasonally, there is a condensation of parasitized hosts towards the end of development as unparasitized nymphs emerge earlier. Spatially, infection levels show a downstream decline that may result from upstream dispersal by infected hosts or differential parasite survivorship at different elevations.     

A laboratory study of predator-prey interactions of stoneflies and mayflies

SUMMARY. 1. Reactions of individual predatory stoneflies to contact by individual mayfly prey, and vice versa, were studied in the laboratory using two sets of species, one from a stream in southern Ontario. Canada, and the other from a stream in North Wales. 2. Based largely on information received via their antennae, the stoneflies were capable of distinguishing between conspecifics and mayfly prey and between prey species. A clear preference for certain prey species was evident but less preferred species were made more attractive when damaged or when the stoneflies were very hungry. Detection of conspecifics by stoneflies was confused in the presence of prey or the local smell of prey body fluids and this led to interference during hunting and feeding at high predator densities. 3. Some mayfly species were capable of distinguishing between conspecifics. other mayflies and predators. The cerci were important in intra- and interspecific contact encounters between mayflies and were used in a fencing fashion to establish a minimum distance between nymphs. However, the cerci did not appear to play an anti-predation role as stoneflies captured and ate mayflies with and without cerci in equal numbers. 4. Reactions to conspecifics by the Palaearetie mayflies Rhithrogena semicolorata (Curt.) and Baetis rhodani (Pictet) were similar to those of the ecologically-equivalent Nearctic species Stenonema vicarium (Walker) and Baetis vagans McDunnough. 5. In encounters between mayflies, intra- and interspecific responses were similar. However, the responses of some mayfly species to conspeeifics were modified in the physical presence of a predator or the water from its holding tank. Different mayfly species made specific responses to predaeeous stoneflies. Responses to a herbivorous stonefly were either similar to conspeeific encounters or were intermediate between them and encounters with predators. 6. Experiments to reveal the exact nature of the stonefly stimulus detected by the mayfly nymphs strongly suggested that there were two components to the stimulus. One was physical contact by the predator's antennae, the other was chemical in nature and could be detected only over short distances (up to a few mm). Independently each could elicit a response, but combined their effect appeared to be synergistic. 7. Different mayfly species responded differently to the stonefly Phasganophora capitata (Pictet) and this sequence of sensitivity was mirrored by the predator's choice of prey species—the most sensitive to the predator being the favoured prey and the less sensitive the least favoured. The significance of these findings to optimal foraging theory, as applied to tactile predators, is discussed.