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The effect of reservoir construction on medically important parasites is well known worldwide but lacks information in Vietnam. With 385 active hydropower plants and numerous water reservoirs, Vietnam provides an ideal setting for studying this issue. This study investigated trematode infection in snail first intermediate hosts from three hydropower reservoirs: Hoa Binh, Son La, and Thac Ba. In total, 25,299 snails representing 16 species were examined, with 959 individuals (8 species) shedding identifiable cercariae. Infection prevalence was highest in thiarid snails (5.4%-15.4%), followed by bithynid snails (2.9%-5.8%). Other snail species showed infection prevalence ranging from 0.3% to 2.9%. Infection prevalence varied significantly across regions, with the highest prevalence in Son La, followed by Hoa Binh and Thac Ba reservoirs. However, no significant differences were observed between snails collected from reservoirs versus canals and paddy fields. Morphological identification resulted in nine cercarial morphotypes, with pleurolophocercaria, xiphidiocercariae, and echinostome being the most common types, accounting for 89.2% of all cercarial infections. Echinostome cercariae were found in seven snail species, while the other cercarial morphotypes were shed by two to five species. Gabbia fuchsiana , Parafossarulus manchouricus , and Melanoides tuberculata were the most common hosts, each harboring five cercarial morphotypes, while Radix auricularia only released echinostome type. In conclusion, our findings highlight the endemic presence of trematodes in hydropower reservoirs and emphasize the need to consider the human-environment interaction around these reservoirs for a better understanding of disease transmission risks.

Schistosomiasis is a parasitic disease affecting more than 250 million people worldwide. Schistosomes infect humans by cercariae penetrating the skin in a freshwater environment. Findings obtained more than 100 years prior showed that miracidium develops into cercaria in freshwater snails, though detailed development dynamics have not been elucidated. Although results of histological analyses of development of schistosomes in snails were presented in our previous studies, findings obtained with dynamic imaging have yet to be reported. In the present study, imaging of schistosome infection and dynamics in snails occuring within a short period was performed using fluorescent labeling agents. Labeling of S. mansoni cercariae with carboxyfluorescein succinimidyl ester (CFSE) caused no toxicity, and allowed for monitoring of schistosome dynamics in snails for up to 10 days and release of infective cercariae without fluorescence in 40 days following infection., Competing Interests: Declaration of competing interest The authors have no conflicts of interest to declare., (Copyright © 2024 Elsevier B.V. All rights reserved.)

Background: Combating infectious diseases and halting biodiversity loss are intertwined challenges crucial to ensure global health. Biodiversity can constrain the spread of vector-borne pathogens circulation, necessitating a deeper understanding of ecological mechanisms underlying this pattern. Our study evaluates the relative importance of biodiversity and the abundance of Bulinus truncatus, a major intermediate host for the trematode Schistosoma haematobium on the circulation of this human pathogen at aquatic transmission sites., Methods: We combined mathematical modelling and a molecular based empirical study to specifically assess the effect of co-infections between S. haematobium and other trematodes within their B. truncatus snail hosts; and B. truncatus abundance at transmission sites, on the production of S. haematobium infective cercariae stages released into the aquatic environment., Results: Our modelling approach shows that more competitive trematode species exploiting B. truncatus as an intermediate host at the transmission site level leads to higher co-infection rates within snail hosts, subsequently reducing the production of S. haematobium cercariae. Conversely, an increase in B. truncatus abundance results in lower co-infection rates, and a higher proportion of S. haematobium cercariae released into the environment. Our empirical data from the field support these findings, indicating a significant negative effect of local trematode species richness (P-value = 0.029; AIC = 14.9) and co-infection rates (P-value = 0.02, AIC = 17.4) on the dominance of S. haematobium based on our GLMM models, while B. truncatus abundance positively influences S. haematobium dominance (P-value = 0.047, AIC = 20.1)., Conclusions: Our study highlights the importance of biodiversity in influencing the transmission of S. haematobium through the effect of antagonistic interactions between trematodes within bulinid snail hosts. This effect intensifies when B. truncatus populations are low, promoting co-infections within snails. In line with the One Health concept, our results suggest that maintaining high level of freshwater biodiversity to sustain global trematode diversity at transmission sites can help reducing the circulation of Schistosoma species locally., (© 2024. The Author(s).)

Background: Mali is known to be a schistosomiasis-endemic country with a limited supply of clean water. This has forced many communities to rely on open freshwater bodies for many human-water contact (HWC) activities. However, the relationship between contact with these water systems and the level of schistosome infection is currently receiving limited attention. This study assessed human-water interactions including cercarial emergence pattern and their influences on urinary schistosomiasis transmission in two communities in the Kayes district of Mali., Methods: We carried out a parasitological study first in children in September 2021, then a cross-sectional study of quantitative observations of human-water contact activities in the population, and finally a study of snail infectivity at contact points in September 2022. The study took place in two communities, Fangouné Bamanan and Diakalèl in the Kayes region of western Mali. The chronobiological study focused on cercarial release from naturally infected snails. Released cercariae were molecularly genotyped by targeting the cox1 region, and the ITS and 18S ribosmal DNA gene (18S rDNA) regions of the DNA. Links between sociodemographic parameters, human water-contact points and hematuria were established using multivariate statistical analysis or the logistic regression model., Results: The main factor predisposing the 97 participants to water contact was domestic activity (62.9%). Of the 378 snails collected at 14 sampling sites, 27 (7.1%) excreted schistosome cercariae, with 15.0% (19/126) at Fangouné Bamanan and 3.3% (8/252) at Diakalel. The release of Schistosoma cercariae shows three different patterns in Fangouné Bamanan: (i) an early release peak (6:00-8:00 AM), (ii) a mid-day release peak (10:00 AM-12:00 PM) and (iii) a double peak: (6:00-8:00 AM) and (6:00-8:00 PM) cercariae release; and two release patterns in Diakalel: early release (6:00-8:00 AM) and (ii) mid-day release (12:00-2:00 PM). All cercariae released during early diurnal (6:00-8:00 AM) or nocturnal emission patterns (6:00-8:00 PM) were hybrids parasite having an cox1 S. bovis or S. curassoni associated with an ITS and 18S rDNA of S. haematobium while the cercariae released during diurnal, or mid-day patterns (8:00 AM-6:00 PM) were pure S. haematobium., Conclusions: Our study showed that domestic activity is the main source of exposure in the Kayes region. Two and three cercariae emission patterns were observed at Diakalel and Fangouné Bamanan respectively. These results suggest that the parasite adapts to the human-water contact period in order to increase its infectivity., (© 2024. The Author(s).)

Background: Schistosomiasis is a parasitic disease which is spread through skin contact with water containing Schistosoma cercariae. Drug treatment has been the main control method, but it does not prevent reinfection. The use of soap can be a complementary measure to reduce transmission. Therefore, this study investigates the quantitative effect of different soaps on the mortality of Schistosoma mansoni cercariae., Methodology: Four soaps including two powder soaps (Kleesoft and Omo) and two bar soaps (B29 and Rungu) which are used in a schistosomiasis-endemic Tanzanian village were studied. S. mansoni cercariae were exposed to powder soaps of 0 (control), 10, 50, 75, 100 and 1000 mg/L and to bar soaps of 0 (control), 100, 500 and 1000 mg/L. The highest concentration of 1000 mg/L was selected based on the laboratory-estimated average soap concentration during handwashing. Cercariae were observed under a microscope after 0, 5, 15, 30, 45 and 60 minutes of exposure to determine their survival., Conclusions: All four soaps can kill S. mansoni cercariae and this lethal effect was related to soap concentration and exposure time. At the highest concentration of 1000 mg/L, all cercariae were dead at 5 minutes post-exposure with two powder soaps and Rungu, while 100% cercarial death was achieved between 5 minutes to 15 minutes for B29. Almost all cercariae survived after being exposed to 10 mg/L powder soaps and 100 mg/L bar soaps for 60 minutes. Powder soaps were more lethal than bar soaps. Considering the widely varying concentrations of soap during real-world hygiene activities and the necessity for a very high soap concentration to eliminate all cercariae in a short 5-minute exposure, providing the efficacy of soap in preventing schistosomiasis becomes challenging. Future studies should investigate whether soap can influence alternative mechanisms such as making cercariae unable to penetrate the skin, thereby providing protection., Competing Interests: The authors have declared that no competing interests exist., (Copyright: © 2024 Zhang et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.)

Objective: To compare the external morphological characteristics and movement patterns between Schistosoma japonicum and S. sinensis cercariae., Methods: S. japonicum and S. sinensis cercariae were heat-fixed, and well-extended cercariae, of 50 each species, were randomly selected for measurement of body length, body width, tail stem length, and tail fork length. The external morphological characteristics of S. japonicum and S. sinensis cercariae were compared. In addition, S. japonicum -infected Oncomelania snails and S. sinensis -infected Tricula snails were observed under a microscope and the movement patterns of S. japonicum and S. sinensis cercariae were compared., Results: The mean body length, body width, tail stem length, and tail fork length were (0.16 ± 0.01), (0.05 ± 0.01), (0.14 ± 0.01) mm and (0.06 ± 0.01) mm for S. japonicum cercariae, and (0.13 ± 0.01), (0.05 ± 0.01), (0.13 ± 0.01) mm and (0.06 ± 0.01) mm for S. sinensis cercariae, respectively, and there were significant differences in terms of cercaria body length ( t = 14.583, P < 0.05) and tail stem length ( t = 3.861, P < 0.05), while no significant differences were seen in terms of body width ( t = 0.896, P > 0.05) or tail fork length ( t = -0.454, P > 0.05). Microscopy revealed that the tails of both S. japonicum and S. sinensis cercariae swung from side to side and there was no significant difference in their movement pattern., Conclusions: S. sinensis and S. japonicum cercariae share highly similar external external morphological characteristics and movement patterns.

Avian schistosomes are snail-borne trematode parasites ( Trichobilharzia spp.) that can cause a nasty skin rash in humans when their cercariae mistake us for their normal bird hosts. We sought to investigate drivers of the spatial distribution of Trichobilharzia cercaria abundance throughout Northern Michigan lakes. For 38 sites on 16 lakes, we assessed several dozen potential environmental predictors that we hypothesized might have direct or indirect effects on overall cercaria abundance, based on known relationships between abiotic and biotic factors in wetland ecosystems. We included variables quantifying local densities of intermediate hosts, temperature, periphyton growth rates, human land use and hydrology. We also measured daily abundance of schistosome cercariae in the water over a 5-week period, supported by community scientists who collected and preserved filtered water samples for qPCR. The strongest predictor of cercaria abundance was Lymnaea host snail density. Lymnaea density was higher in deeper lakes and at sites with more deciduous tree cover, consistent with their association with cool temperature habitats. Contrary to past studies of human schistosomes, we also found a significant negative relationship between cercaria abundance and submerged aquatic vegetation, possibly due to vegetation blocking cercaria movement from offshore snail beds. If future work shows that these effects are indeed causal, then these results suggest possible new approaches to managing swimmer's itch risk in northern MI lakes, such as modifying tree cover and shallow-water vegetation at local sites.

Background: Trematode infections of the genus Schistosoma can induce physiological and behavioral changes in intermediate snail hosts. This is because the parasite consumes essential resources necessary for the host's survival, prompting hosts to adapt their behavior to maintain some level of fitness before parasite-induced mortality occurs., Methods: In this study, the reproductive and biochemical parameters of Biomphalaria alexandrina and Bulinus truncatus were examined during the cercareal shedding stage of infection with Schistosoma mansoni and Schistosoma haematobium, respectively, compared with controls., Results: The study revealed an infection rate of 34.7% for S. mansoni and 30.4% for S. haematobium. In B. alexandrina infected with S. mansoni, a survival rate of 65.2% was recorded, along with a mean prepatent period of 30.3 ± 1.41 days, a mean shedding duration of 14.2 ± 0.16 days, and a mean lifespan of 44.1 ± 0.24 days. Meanwhile, in B. truncatus infected with S. haematobium, a survival rate of 56.4% was observed, with a mean prepatent period of 44.3 ± 1.41 days, a mean shedding duration of 22.6 ± 2.7 days, and a mean lifespan of 66.9 ± 1.6 days. Feeding increased in both infected species of snails, while the net reproductive rate (Ro) of the infected snails decreased. Total antioxidant (TAO) and lipid peroxidation activity increased in the two infected snail species during shedding, while Glutathione-S-transferase levels decreased. Lipid peroxidase activity and nitrogen oxide levels significantly decreased in infected B. alexandrina and increased in infected Bulinus. Steroid hormone levels were elevated in infected Biomphalaria, whereas they were reduced in infected Bulinus. Comet assay parameters showed an increase in the two infected genera after infection compared to control snails, indicating genotoxic damage and histopathological damage was observed., Conclusions: These findings demonstrate that infection with larva species diverse biochemical, hormonal, genotoxic, and histopathological changes in the tissues responsible for fecundity and reproduction in B. alexandrina and B. truncates comparing with controls., (© 2024. The Author(s).)

The lifecycle of schistosomes must be continuously maintained to clarify and understand this parasite in various aspects in laboratory settings. In the previous studies by other researchers, preservation of schistosome larvae or eggs was attempted by freezing with liquid nitrogen or organic chemicals, but frozen schistosomes were substantially impaired. The present study was conducted to determine whether schistosome eggs can be preserved under a non-frozen condition. The results showed that Schistosoma mansoni eggs could be maintained in phosphate-buffered saline at 4 °C, with a high level of infectivity of miracidia to freshwater snails thereafter. Furthermore, the egg hatchability was maintained for up to 12 weeks with weekly exchanges of the medium. The cercariae derived from snails infected with miracidia from preserved eggs were highly infective to mice. This simple schistosome egg preservation method allow researchers to maintain the schistosome lifecycle without freezing or other special procedures., Competing Interests: Declaration of competing interest The authors have no conflicts of interest to declare., (Copyright © 2024 Elsevier B.V. All rights reserved.)

Global warming is likely to lengthen the seasonal duration of larval release by parasites. We exposed freshwater mussel hosts, Anodonta anatina , from 2 high-latitude populations to high, intermediate and low temperatures throughout the annual cercarial shedding period of the sympatric trematodes Rhipidocotyle fennica and R. campanula , sharing the same transmission pathway. At the individual host level, under warmer conditions, the timing of the cercarial release in both parasite species shifted towards seasonally earlier period while its duration did not change. At the host population level, evidence for the lengthening of larvae shedding period with warming was found for R. fennica . R. campanula started the cercarial release seasonally clearly earlier, and at a lower temperature, than R. fennica . Furthermore, the proportion of mussels shedding cercariae increased, while day-degrees required to start the cercariae shedding decreased in high-temperature treatment in R. fennica . In R. campanula these effects were not found, suggesting that warming can benefit more R. fennica . These results do not completely support the view that climate warming would invariably increase the seasonal duration of larval shedding by parasites, but emphasizes species-specific differences in temperature-dependence and in seasonality of cercarial release.

Cercarial activity and survival are crucial traits for the transmission of trematodes. Temperature is particularly important, as faster depletion of limited cercarial energy reserves occurs at high temperatures. Seasonal climate conditions in high latitude regions may be challenging to complete trematode life cycle during the 6-month ice-free period, but temperature effects on the activity and survival of freshwater cercariae have not been previously identified. After experimentally simulating natural subarctic conditions during warmer and colder months (13 and 6°C), a statistical approach identifying changes in the tendency of cercarial activity loss and mortality data was used to detect differences in three trematode genera, represented by four taxa (Diplostomum spp., Apatemon spp., small- and large-sized Plagiorchis spp.). A strong temperature-dependent response was identified in both activity loss and mortality in all taxa, with Diplostomum spp. cercariae showing the most gradual changes compared to other taxa. Furthermore, whilst activity loss and mortality dynamics could not be divided into ‘fish- vs invertebrate-infecting cercariae’ groups, the detected taxa-specific responses in relation to life-history traits indicate the swimming behaviour of cercariae and energy allocation among larvae individuals as the main drivers. Cercariae exploit the short transmission window that allows a stable continuance of trematodes’ life cycles in high-latitude freshwater ecosystems.

Seawater intrusion associated with decreasing groundwater levels and rising seawater levels may affect freshwater species and their parasites. While brackish water certainly impacts freshwater systems globally, its impact on disease transmission is largely unknown. This study examined the effect of artificial seawater on host-parasite interactions using a freshwater snail host, Biomphalaria alexandrina, and the human trematode parasite Schistosoma mansoni. To evaluate the impact of increasing salinity on disease transmission four variables were analyzed: snail survival, snail reproduction, infection prevalence, and the survival of the parasite infective stage (cercariae). We found a decrease in snail survival, snail egg mass production, and snail infection prevalence as salinity increases. However, cercarial survival peaked at an intermediate salinity value. Our results suggest that seawater intrusion into freshwaters has the potential to decrease schistosome transmission to humans., Competing Interests: The authors have declared that no competing interests exist.

To predict global warming impacts on parasitism, we should describe the thermal tolerance of all players in host-parasite systems. Complex life-cycle parasites such as trematodes are of particular interest since they can drive complex ecological changes. This study evaluates the net response to temperature of the infective larval stage of Himasthla elongata, a parasite inhabiting the southwestern Baltic Sea. The thermal sensitivity of (i) the infected and uninfected first intermediate host (Littorina littorea) and (ii) the cercarial emergence, survival, self-propelling, encystment, and infection capacity to the second intermediate host (Mytilus edulis sensu lato) were examined. We found that infection by the trematode rendered the gastropod more susceptible to elevated temperatures representing warm summer events in the region. At 22 °C, cercarial emergence and infectivity were at their optimum while cercarial survival was shortened, narrowing the time window for successful mussel infection. Faster out-of-host encystment occurred at increasing temperatures. After correcting the cercarial emergence and infectivity for the temperature-specific gastropod survival, we found that warming induces net adverse effects on the trematode transmission to the bivalve host. The findings suggest that gastropod and cercariae mortality, as a tradeoff for the emergence and infectivity, will hamper the possibility for trematodes to flourish in a warming ocean., (© 2022. The Author(s).)

Parasites remainunderstudied members of most ecosystems, especially free-living infectious stages, such as the aquatic cercariae of trematodes (flatworms). Recent studies are shedding more light on their roles, particularly as prey for a diverse array of aquatic predators, but the possible fates of cercariae remain unclear. While this is critical to elucidate because cercariae represent a large potential source of energy and nutrients, determining the fate of cercariae-derived organic matter involves many logistical challenges. Previous studies utilized elemental and stable isotope analysis when examining host-parasite interactions, but none has used such approaches to track the movement of cercariae biomass within food webs. Here we report that Plagiorchis sp. cercariae were effectively labelled with 13 C by introducing this compound in the food of their snail host. We then added 13 C-labelled cercariae as a potential food source to experimental mesocosms containing a simplified model freshwater food web represented by diving beetles (Dytiscidae sp.), dragonfly larvae (Leucorrhinia intacta), oligochaete worms (Lumbriculus variegatus), and a zooplankton community dominated by Daphnia pulex. The oligochaetes had the highest ratio of 13 C to 12 C, suggesting benthic detritivores are substantial, but previously unrecognized, consumers of cercariae biomass. In an experiment where L. variegatus were fed mass equivalents of dead D. pulex or cercariae, growth was greater with the latter diet, supporting the importance of cercariae as food source for benthic organisms. Given the substantial cercariae biomass possible in natural settings, understanding their contributions to energy flow and nutrient cycling is important, along with developing methods to do so.

Life cycle stages, including daughter sporocysts, cercariae, and metacercariae, of Parvatrema duboisi (Dollfus, 1923) Bartoli, 1974 (Digenea: Gymnophallidae) have been found in the Manila clam Ruditapes philippinarum from Aphaedo (Island), Shinan-gun, Jeollanam-do, Korea. The daughter sporocysts were elongated sac-like and 307-570 (av. 395) μm long and 101-213 (av. 157) μm wide. Most of the daughter sporocysts contained 15-20 furcocercous cercariae each. The cercariae measured 112-146 (av. 134) μm in total length and 35-46 (av. 40) μm in width, with 69-92 (av. 85) μm long body and 39-54 (av. 49) μm long tail. The metacercariae were 210-250 (av. 231) μm in length and 170-195 (av. 185) μm in width, and characterized by having a large oral sucker, genital pore some distance anterior to the ventral sucker, no ventral pit, and 1 compact or slightly lobed vitellarium, strongly suggesting P. duboisi. The metacercariae were experimentally infected to ICR mice, and adults were recovered at day 7 post-infection. The adult flukes were morphologically similar to the metacercariae except in the presence of up to 20 eggs in the uterus. The daughter sporocysts and metacercariae were molecularly (ITS1-5.8S rDNA-ITS2) analyzed to confirm the species, and the results showed 99.8-99.9% identity with P. duboisi reported from Kyushu, Japan and Gochang, Korea. These results confirmed the presence of various life cycle stages of P. duboisi in the Manila clam, R. philippinarum, playing the role of the first as well as the second intermediate host, on Aphae-do (Island), Shinan-gun, Korea.

Background: Leeches (Hirudinida) play a significant role as intermediate hosts in the circulation of trematodes in the aquatic environment. However, species richness and the molecular diversity and phylogeny of larval stages of strigeid trematodes (tetracotyle) occurring in this group of aquatic invertebrates remain poorly understood. Here, we report our use of recently obtained sequences of several molecular markers to analyse some aspects of the ecology, taxonomy and phylogeny of the genera Australapatemon and Cotylurus, which utilise leeches as intermediate hosts., Methods: From April 2017 to September 2018, 153 leeches were collected from several sampling stations in small rivers with slow-flowing waters and related drainage canals located in three regions of Poland. The distinctive forms of tetracotyle metacercariae collected from leeches supplemented with adult Strigeidae specimens sampled from a wide range of water birds were analysed using the 28S rDNA partial gene, the second internal transcribed spacer region (ITS2) region and the cytochrome c oxidase (COI) fragment., Results: Among investigated leeches, metacercariae of the tetracotyle type were detected in the parenchyma and musculature of 62 specimens (prevalence 40.5%) with a mean intensity reaching 19.9 individuals. The taxonomic generic affiliation of metacercariae derived from the leeches revealed the occurrence of two strigeid genera: Australapatemon Sudarikov, 1959 and Cotylurus Szidat, 1928. Phylogenetic reconstructions based on the partial 28S rRNA gene, ITS2 region and partial COI gene confirmed the separation of the Australapatemon and Cotylurus clades. Taking currently available molecular data and our results into consideration, recently sequenced tetracotyle of Australapatemon represents most probably Au. minor; however, unclear phylogenetic relationships between Au. burti and Au. minor reduce the reliability of this conclusion. On the other hand, on the basis of the obtained sequences, supplemented with previously published data, the metacercariae of Cotylurus detected in leeches were identified as two species: C. strigeoides Dubois, 1958 and C. syrius Dubois, 1934. This is the first record of C. syrius from the intermediate host., Conclusions: The results of this study suggest the separation of ecological niches and life cycles between C. cornutus (Rudolphi, 1808) and C. strigeoides/C. syrius, with potential serious evolutionary consequences for a wide range of host-parasite relationships. Moreover, phylogenetic analyses corroborated the polyphyletic character of C. syrius, the unclear status of C. cornutus and the separate position of Cotylurus raabei Bezubik, 1958 within Cotylurus. The data demonstrate the inconsistent taxonomic status of the sequenced tetracotyle of Australapatemon, resulting, in our opinion, from the limited availability of fully reliable, comparative sequences of related taxa in GenBank.

Australapatemon spp. are cosmopolitan trematodes that infect freshwater snails, aquatic leeches, and birds. Despite their broad geographic distribution, relatively little is known about interactions between Australapatemon spp. and their leech hosts, particularly under experimental conditions and in natural settings. We used experimental exposures to determine how Australapatemon burti cercariae dosage (number administered to leech hosts, Erpobdella microstoma) affected infection success (fraction to encyst as metacercariae), infection abundance, host survival, and host size over the 100 days following exposure. Interestingly, infection success was strongly density-dependent, such that there were no differences in metacercariae load even among hosts exposed to a 30-fold difference in cercariae. This relationship suggests that local processes (e.g., resource availability, interference competition, or host defenses) may play a strong role in parasite transmission. Our results also indicated that metacercariae did not become evident until ~4 weeks post exposure, with average load climbing until approximately 13 weeks. There was no evidence of metacercariae death or clearance over the census period. Parasite exposure had no detectable effects on leech size or survival, even with nearly 1,000 cercariae. Complementary surveys of leeches in California revealed that 11 of 14 ponds supported infection by A. burti (based on morphology and molecular sequencing), with an average prevalence of 32% and similar metacercariae intensity as in our experimental exposures. The extended development time and extreme density dependence of A. burti has implications for studying naturally occurring host populations, for which detected infections may represent only a fraction of cercariae to which animals have been exposed. Future investigation of these underlying mechanisms would be benefical in understanding host-parasite relationships., (Published by Elsevier Inc.)

Direct consumption on free-living cercariae stages of trematodes by non-host organisms interferes with trematode transmission and leads to reduced infections in the next suitable hosts. Consumer functional responses provide a useful tool to examine relationships between consumption rates and ecologically relevant prey densities, whilst also accounting for abiotic factors that likely influence consumption rates. We investigated how temperature influences the consumer functional response of the amphipod Gammarus lacustris towards the cercariae of three freshwater trematodes (Diplostomum, Apatemon and Trichobilharzia). Amphipods displayed different functional responses towards the parasites, with Type II responses for Diplostomum and Type I responses for Apatemon prey. Temperature did not alter the consumption rate of the amphipod predator. Trichobilharzia was likely consumed at similar proportions as Diplostomum; however, this could not be fully evaluated due to low replication. Whilst Type II responses of invertebrate predators are common to various invertebrate prey types, this is the first time a non-filter feeding predator has been shown to exhibit Type I response towards cercarial prey. The prey-specific consumption patterns of amphipods were related to cercarial distribution in the water column rather than to the size of cercariae or temperature influence. The substantial energy flow into food webs by non-host consumer organisms highlights the importance of understanding the mechanisms that modulate functional responses and direct predation in the context of parasitic organisms.

Cercariae invasion of the human skin is the first step in schistosome infection. Proteases play key roles in this process. However, little is known about the related hydrolytic enzymes in Schistosoma japonicum. Here, we investigated the biochemical features, tissue distribution and biological roles of a cathepsin B cysteine protease, SjCB2, in the invasion process of S. japonicum cercariae. Enzyme activity analysis revealed that recombinant SjCB2 is a typical cysteine protease with optimum temperature and pH for activity at 37°C and 4.0, respectively, and can be totally inhibited by the cysteine protease inhibitor E-64. Immunoblotting showed that both the zymogen (50 kDa) and mature enzyme (30.5 kDa) forms of SjCB2 are expressed in the cercariae. It was observed that SjCB2 localized predominantly in the acetabular glands and their ducts of cercariae, suggesting that the protease could be released during the invasion process. The protease degraded collagen, elastin, keratin, fibronectin, immunoglobulin (A, G and M) and complement C3, protein components of the dermis and immune system. In addition, proteomic analysis demonstrated that SjCB2 can degrade the human epidermis. Furthermore, it was showed that anti-rSjCB2 IgG significantly reduced (22.94%) the ability of the cercariae to invade the skin. The cysteine protease, SjCB2, located in the acetabular glands and their ducts of S. japonicum cercariae. We propose that SjCB2 facilitates skin invasion by degrading the major proteins of the epidermis and dermis. However, this cysteine protease may play additional roles in host-parasite interaction by degrading immunoglobins and complement protein., Competing Interests: The authors have declared that no competing interests exist.

Trematode transmission in aquatic habitats from molluscan intermediate host to vertebrate or invertebrate target host is typically undertaken by a free-living stage known as cercariae. Active locomotion by cercariae is a key aspect of the transmission process with the swimming speed potentially contributing to infection success. Individual cercarial species swim at different speeds but the significance of this to infection potential has not been determined. This study, using data from the scientific literature, investigates the role of swimming speed in relation to cercarial morphology, host-searching strategies and target host species. Larger cercariae swim faster than smaller ones with tail length being the principal factor controlling locomotion rates. Different cercarial morphotypes swim at different speeds, in particular, furcocercariae, with the exception of the schistosomes, being faster swimmers than mono-tailed cercariae. Host-searching behaviour has a significant influence on swimming speeds with 'active-searching' strategies swimming slower than those adopting 'active-waiting' or 'prey mimcry' strategies. Vertebrate-infecting cercariae swim faster than those infecting invertebrates with species targeting fish demonstrating the highest locomotion rates and those targeting arthropods the slowest speeds. The adaptions of individual cercarial swimming speeds to biological variables and their interactions with the physical processes of aquatic habitats are discussed.

Parasitic castration of bivalves by trematodes is common, and may significantly reduce the reproductive capacity of ecologically important species. Understanding the intensity of infection is desirable, as it can indicate the time that has passed since infection, and influence the host's physiological and reproductive response. In addition, it is useful to know the developmental stage of the trematode, to understand trematode population trends and reproductive success. However, most existing methods (e.g. visually estimating the degree of infection) to assess intensity are approximate only and not reproducible. Here, we present a method to accurately quantify the percentage of bivalve gonad filled with digenean trematode tissue, based on small squashes of gonad tissue rapidly photographed under light microscopy. A maximum of 15 photographs is required to determine the percentage of the whole gonad occupied by trematodes with a minimum of 90% confidence, with smaller mussels requiring fewer. In addition, the stage of trematode infection can be assessed because full sporocysts, spent sporocysts and free cercariae are clearly distinguishable. Although variation exists in the distribution of trematodes in gonad tissue, and thus in the estimate of percentage of the gonad filled with trematodes, this method represents a marked improvement on current coarse assessments of infection which typically focus on binary presence/absence measures. This technique can be used to facilitate a more sophisticated understanding of host-parasite interactions in bivalves, and can inform the conservation and reproductive biology of environmentally crucial species.

Background: Cercarial dermatitis (swimmer's itch) caused by bird schistosome cercariae, released from intermediate host snails, is a common disorder also at higher latitudes. Several cases were observed in the artificial Danish freshwater Ringen Lake frequently used by the public for recreational purposes. The lake may serve as a model system when establishing a risk analysis for this zoonotic disease. In order to explain high risk periods we determined infection levels of intermediate host snails from early spring to late summer (March, June and August) and elucidated the effect of temperature and light on parasite shedding, behavior and life span., Results: Field studies revealed no shedding snails in March and June but in late summer the prevalence of Trichobilharzia szidati infection (in a sample of 226 pulmonate Lymnaea stagnalis snails) reached 10%. When investigated under laboratory conditions the cercarial shedding rate (number of cercariae shed per snail per day) was positively correlated to temperature raising from a mean of 3000 (SD 4000) at 7 °C to a mean of 44,000 (SD 30,000) at 27 °C). The cercarial life span was inversely correlated to temperature but the parasites remained active for up to 60 h at 20 °C indicating accumulation of cercariae in the lake during summer periods. Cercariae exhibited positive phototaxy suggesting a higher pathogen concentration in surface water of the lake during daytime when the public visits the lake., Conclusion: The only causative agent of cercarial dermatitis in Ringen Lake detected was T. szidati. The infection risk associated with aquatic activities is low during spring and early summer (March-June). In late summer the risk of infection is high since the release, behavior and life span of the infective parasite larvae have optimal conditions.

Background: Schistosomiasis is responsible for the second highest burden of disease among neglected tropical diseases globally, with over 90 percent of cases occurring in African regions where drugs to treat the disease are only sporadically available. Additionally, human re-infection after treatment can be a problem where there are high numbers of infected snails in the environment. Recent experiments indicate that aquatic factors, including plants, nutrients, or predators, can influence snail abundance and parasite production within infected snails, both components of human risk. This study investigated how snail host abundance and release of cercariae (the free swimming stage infective to humans) varies at water access sites in an endemic region in Senegal, a setting where human schistosomiasis prevalence is among the highest globally., Methods/principal Findings: We collected snail intermediate hosts at 15 random points stratified by three habitat types at 36 water access sites, and counted cercarial production by each snail after transfer to the laboratory on the same day. We found that aquatic vegetation was positively associated with per-capita cercarial release by snails, probably because macrophytes harbor periphyton resources that snails feed upon, and well-fed snails tend to produce more parasites. In contrast, the abundance of aquatic macroinvertebrate snail predators was negatively associated with per-capita cercarial release by snails, probably because of several potential sublethal effects on snails or snail infection, despite a positive association between snail predators and total snail numbers at a site, possibly due to shared habitat usage or prey tracking by the predators. Thus, complex bottom-up and top-down ecological effects in this region plausibly influence the snail shedding rate and thus, total local density of schistosome cercariae., Conclusions/significance: Our study suggests that aquatic macrophytes and snail predators can influence per-capita cercarial production and total abundance of snails. Thus, snail control efforts might benefit by targeting specific snail habitats where parasite production is greatest. In conclusion, a better understanding of top-down and bottom-up ecological factors that regulate densities of cercarial release by snails, rather than solely snail densities or snail infection prevalence, might facilitate improved schistosomiasis control., Competing Interests: The authors have declared that no competing interests exist.

Opisthorchis viverrini is a fish-borne zoonotic trematode that causes significant public health problems in Southeast Asia. Its life cycle requires Bithynia snails as the first intermediate hosts, fish, and human and/or carnivore hosts. This study assessed impacts of land use practice for rice cultivation and seasonality on the transmission dynamics of O. viverrini in Bithynia siamensis goniomphalos in rice paddy field habitats. The transmission of O. viverrini cercariae in B. s. goniomphalos was monitored at monthly intervals over a 4-year period from January 2010 to December 2013. From a total of 59,727 snails examined by standard cercarial shedding, the prevalence of O. viverrini was 0.7% (range, 0.0-4.1%). The prevalence of O. viverrini infection in B. s. goniomphalos varied with the amount of rainfall, with peaks of infection occurring in the cool-dry season, that is, after each rainy season. A shift of peak prevalence from cool-dry to hot-dry season observed in 2013 was associated with the increase in preceding water irrigation to support the production of second annual rice crop. Significant positive correlations were found between the prevalence and intensity of cercarial infection and the size of snails. Our results revealed substantial variation between years so that to have a clear understanding of the population dynamics of this complex system, studies should be conducted over an extended period (> 1 year). Results from this study highlight that water irrigation schemes in rice paddy cultivation and seasonality have a significant effect on the prevalence of O. viverrini in B. s. goniomphalos .

Infection with trematodes produces physiological and behavioural changes in intermediate snail hosts. One response to infection is parasitic castration, in which energy required for reproduction of the host is thought to be redirected to promote development and multiplication of the parasite. This study investigated some reproductive and biochemical parameters in the nervous (CNS) and ovotestis (OT) tissues of Biomphalaria alexandrina during the course of Schistosoma mansoni infection. Antioxidant and oxidative stress parameters including catalase (CAT), nitric oxide (NO) and lipid peroxidation (MDA) were measured. Levels of steroid hormones, including testosterone, progesterone and estradiol, were also assessed. Finally, flow cytometry was used to compare measures of apoptosis between control snails and those shedding cercariae by examining mitochondrial membrane potential with the stain 5,5',6,6'-tetrachloro-1,1',3,3'-tetraethylbenzimi-dazolylcarbocyanine iodide (JC-1) and poly(ADP-ribose) polymerase (PARP). Infection with S. mansoni caused a 47.7% reduction in the net reproductive rate (R o ) of B. alexandrina. CAT activity was increased in the CNS at 21 days post infection (dpi) but by 28 dpi it was reduced below control values. Also, CAT activity increased significantly in the OT at 14, 21 and 28 dpi. In CNS tissues, NO levels were reduced at 7 dpi, increased at 14 and 21 dpi, and reduced again at 28 dpi. The overall level of lipid peroxidation gradually increased during the course of infection to reach its highest levels at 28 dpi. Steroid hormone measurements showed that concentrations of testosterone and estradiol were reduced in the CNS tissues at 28 dpi, while those of progesterone were slightly increased in the CNS and OT tissues. The percentage of cells that positively stained with JC-1was significantly increased in CNS and OT tissues of infected snails while the percentage of cells positively stained with PARP was decreased compared to controls. Together, these findings indicate that infection initiates diverse biochemical and hormonal changes leading to loss of cells responsible for egg laying and reproduction in B. alexandrina., Competing Interests: Declaration of competing interest The authors declare that they have no competing interests., (Copyright © 2020 Elsevier Inc. All rights reserved.)

Background: Urogenital schistosomiasis, caused by infection with Schistosoma haematobium, is endemic in Niger but complicated by the presence of Schistosoma bovis, Schistosoma curassoni and S. haematobium group hybrids along with various Bulinus snail intermediate host species. Establishing the schistosomes and snails involved in transmission aids disease surveillance whilst providing insights into snail-schistosome interactions/compatibilities and biology., Methods: Infected Bulinus spp. were collected from 16 villages north and south of the Niamey region, Niger, between 2011 and 2015. From each Bulinus spp., 20-52 cercariae shed were analysed using microsatellite markers and a subset identified using the mitochondrial (mt) cox1 and nuclear ITS1 + 2 and 18S DNA regions. Infected Bulinus spp. were identified using both morphological and molecular analysis (partial mt cox1 region)., Results: A total of 87 infected Bulinus from 24 sites were found, 29 were molecularly confirmed as B. truncatus, three as B. forskalii and four as B. globosus. The remaining samples were morphologically identified as B. truncatus (n = 49) and B. forskalii (n = 2). The microsatellite analysis of 1124 cercariae revealed 186 cercarial multilocus genotypes (MLGs). Identical cercarial genotypes were frequently (60%) identified from the same snail (clonal populations from a single miracidia); however, several (40%) of the snails had cercariae of different genotypes (2-10 MLG's) indicating multiple miracidial infections. Fifty-seven of the B. truncatus and all of the B. forskalii and B. globosus were shedding the Bovid schistosome S. bovis. The other B. truncatus were shedding the human schistosomes, S. haematobium (n = 6) and the S. haematobium group hybrids (n = 13). Two B. truncatus had co-infections with S. haematobium and S. haematobium group hybrids whilst no co-infections with S. bovis were observed., Conclusions: This study has advanced our understanding of human and bovid schistosomiasis transmission in the Niger River Valley region. Human Schistosoma species/forms (S. haematobium and S. haematobium hybrids) were found transmitted only in five villages whereas those causing veterinary schistosomiasis (S. bovis), were found in most villages. Bulinus truncatus was most abundant, transmitting all Schistosoma species, while the less abundant B. forskalii and B. globosus, only transmitted S. bovis. Our data suggest that species-specific biological traits may exist in relation to co-infections, snail-schistosome compatibility and intramolluscan schistosome development.

Many temperate freshwater habitats are at risk for contamination by run-off associated with the application of road de-icing salts. Elevated salinity can have various detrimental effects on freshwater organisms, including greater susceptibility to infection by parasites and pathogens. However, to better understand the net effects of road salt exposure on host-parasite dynamics, it is necessary to consider the impacts on free-living parasite infectious stages, such as the motile aquatic cercariae of trematodes. Here, we examined the longevity and activity of cercariae from four different freshwater trematodes (Ribeiroia ondatrae, Echinostoma sp., Cephalogonimus sp. and an unidentified strigeid-type) that were exposed to road salt at five different environmentally relevant concentrations (160, 360, 560, 760 and 960 mg/ml of sodium chloride). Exposure to road salt had minimal detrimental effects, with cercariae activity and survival often greatest at intermediate concentrations. Only the cercariae of Cephalogonimus sp. showed reduced longevity at the highest salt concentration, with those of both R. ondatrae and the unidentified strigeid-type exhibiting diminished activity, indicating interspecific variation in response. Importantly, cercariae seem to be relatively unaffected by salt concentrations known to increase infection susceptibility in some of their hosts. More studies will be needed to examine this potential dichotomy in road salt effects between hosts and trematodes, including influences on parasite infectivity.

In a range of trematode species, specific members of the parthenitae colony infecting the molluscan first intermediate host appear specialized for defence against co-infecting species. The evolution of such division of labour requires that co-infection entails fitness costs. Yet, this premise has very rarely been tested in species showing division of labour. Using Himasthla elongata (Himasthlidae) and Renicola roscovita (Renicolidae) infecting periwinkles Littorina littorea as study system, we show that the size of emerged cercariae is markedly reduced in both parasite species when competing over host resources. Cercarial longevity, on the other hand, is negatively influenced by competition only in R. roscovita. Season, which may impact the nutritional state of the host, also affects cercarial size, but only in H. elongata. Hence, our study underlines that cercarial quality is, indeed, compromised by competition, not only in the inferior R. roscovita (no division of labour) but also in the competitively superior H. elongata (division of labour).

Digenean larvae of hermaphroditic generation - cercariae - are known to be polymorphic at genetic and behavioural levels. Cercariae arise as a result of parthenogenetic reproduction of intramolluscan stages, and represent a clone if a snail was infected with a single miracidium. Here we investigated cercarial clones of Himasthla elongata - namely, the infectivity of cercariae with normal (negative) and deviant (positive) photoreaction. In our study, most H. elongata clones showed intraclonal variance in their response to light. The proportion of photopositive cercariae ranged between 0.2% and 60% in different H. elongata clones. Photopositive larvae demonstrated significantly reduced rates of encystment in Mytilus edulis haemolymph in vitro and in young mussels. We discuss the possible mechanisms behind intraclonal variations, such as non-specific genomic rearrangements.

Parasites with complex life cycles can be susceptible to temperature shifts associated with seasonal changes, especially as free-living larvae that depend on a fixed energy reserve to survive outside the host. The life cycle of Schistosoma, a trematode genus containing some species that cause human schistosomiasis, has free-living, aquatic miracidial and cercarial larval stages that swim using cilia or a forked tail, respectively. The small size of these swimmers (150-350 µm) dictates that their propulsion is dominated by viscous forces. Given that viscosity inhibits the swimming ability of small organisms and is inversely correlated with temperature, changes in temperature should affect the ability of free-living larval stages to swim and locate a host. By recording miracidial and cercarial movement of Schistosoma mansoni using a high-speed camera and manipulating temperature and viscosity independently, we assessed the role each factor plays in the swimming mechanics of the parasite. We found a positive effect of temperature and a negative effect of viscosity on miracidial and cercarial speed. Reynolds numbers, which describe the ratio of inertial to viscous forces exerted on an aquatic organism, were <1 across treatments. Q 10 values were <2 when comparing viscosity treatments at 20 °C and 30 °C, further supporting the influence of viscosity on miracidial and cercarial speed. Given that both larval stages have limited energy reserves and infection takes considerable energy, successful transmission depends on both speed and lifespan. We coupled our speed data with mortality measurements across temperatures and discovered that the theoretical maximum distance travelled increased with temperature and decreased with viscosity for both larval stages. Thus, our results suggest that S. mansoni transmission is high during warm times of the year, partly due to improved swimming performance of the free-living larval stages, and that increases in temperature variation associated with climate change might further increase transmission., (Copyright © 2020 Australian Society for Parasitology. Published by Elsevier Ltd. All rights reserved.)

Field investigations in 14 wild watercress beds located in the French region of Limousin, a known endemic area for distomatosis, were performed for three years to determine the distance that cercariae of Fasciola hepatica can reach in water before their encystment on the host plant. Each bed was located on the course of an open drainage furrow, while snails (Galba truncatula) lived upstream around the emergence of a source. Five plant species were collected in early April and examined to find metacercariae. Most cysts were noted on Nasturtium officinale (188 on 48.7 kg of dripped plants), followed by Helosciadium nodiflorum (125 on 33.4 kg). On the other plant species, there were few larvae. Most cercariae encysted on the plants growing in the most upstream part of each bed, usually on the first 50 cm in length. When water in the beds was fast running, the distribution of metacercariae was more limited and their number was fewer than those in the beds fed by a slow flow of water. Cercariae were able to swim or were carried away by the current up to a mean of 5 m in slow-flow waters before encysting; this distance was only 4 m in faster waters. Plants growing on the most upstream section of a watercress bed located in a drainage furrow are the most used by cercariae for their encystment, when snails live around the emergence of a source. The speed of the water current affected the number and distribution of metacercariae in the bed., (© D. Rondelaud et al., published by EDP Sciences, 2020.)

Individual developmental stages of blood fluke Schistosoma mansoni excrete or secrete a different set of molecules. Here we describe optimized protocols for collection of excretory/secretory products (E/S products) from cercariae, schistosomula, adult worms, and eggs. These E/S products are essential for successful parasitism functioning at the host-parasite interface, enabling invasion into the host and contributing to the survival of the parasite by modulation of host physiology and immune responses. Collection of sufficient amounts of E/S products is required for detailed research of these processes.

Schistosomes are obligate helminths responsible for over 218 million cases of human schistosomiasis in 78 countries around the world. Infection occurs when free-swimming cercariae penetrate human skin and initiate developmental progression into parasitic obligate worms that consume red blood cells. Transcriptomic studies of infectious cercariae reveal abundant mRNAs associated with energy metabolism and host invasion. However, the cercaria is mostly transcriptionally quiescent, suggesting that most mRNAs are primed prior to cercarial escape from the snail host. The use of transcriptomics to understand protein expression presumes that transcription and translation are functionally coupled and the cercarial stage has categorically been treated as a single unit for -omic analysis. Per contra, the relationship between transcription and translation in infectious cercariae has not been described. To understand the correlation between transcription and translation in cercariae, we separately measured nascent translation levels in cercarial heads, cercarial tails and in the developing schistosomula, the next stage of its life cycle. The loss of the cercarial tail is essential for the transformation from a cercaria to a schistosomulum. We observed that translation was initially limited and the translation rate accelerated during the first 72-hours after tail loss. When we tested nascent translation in cercarial heads, cercarial tails, whole cercariae, and 4-hour schistosomula, we found that translation is significantly upregulated in the cercarial tail when compared to the cercarial head and that translation was undetectable in heads using immunofluorescent image quantification (p = .0005). These data represent a major shift in how we understand the cercarial stage. The cercarial head is mostly transcriptionally and translationally quiescent while being sufficient for progression into a schistosomulum. In addition, transcription and translation are not linked in Schistosoma mansoni cercaria. Thus, our current conceptual approach of treating the cercaria as a single functional unit for -omic studies may be insufficient to understand cercarial development., Competing Interests: The authors have declared that no competing interests exist.

Schistosomes are parasitic flatworms that infect over 200 million people, causing the neglected tropical disease, schistosomiasis. A single drug, praziquantel, is used to treat schistosome infection. Limitations in mass drug administration programs and the emergence of schistosomiasis in nontropical areas indicate the need for new strategies to prevent infection. It has been known for several decades that rotifers colonizing the schistosome's snail intermediate host produce a water-soluble factor that paralyzes cercariae, the life cycle stage infecting humans. In spite of its potential for preventing infection, the nature of this factor has remained obscure. Here, we report the purification and chemical characterization of Schistosome Paralysis Factor (SPF), a novel tetracyclic alkaloid produced by the rotifer Rotaria rotatoria. We show that this compound paralyzes schistosome cercariae and prevents infection and does so more effectively than analogous compounds. This molecule provides new directions for understanding cercariae motility and new strategies for preventing schistosome infection., Competing Interests: I have read the journal's policy and the authors of this manuscript have the following competing interest: A patent application has been filed by the Wisconsin Alumni Research Foundation and is pending. Use of a Rotifer-Derived Compound and its Analogs for Preventing Schistosomiasis. US Application #: 16/445766, covers the use of this novel molecule and its derivatives in preventing schistosome infection.

An analysis of daily water samples collected from an index site on Big Soos Creek, Washington indicated intra-annual differences in the concentrations of waterborne Nanophyetus salmincola. Waterborne concentrations, quantified as gene copies/L, peaked during the fall (October-November 2016), decreased to very low concentrations over the winter (January-March 2017), and then increased in the spring and throughout the summer. High waterborne concentrations of N. salmincola DNA (2 × 10 6 gene copies/L) corresponded with live N. salmincola cercariae (mean = 3 cercariae/L) that were detected in companion water samples. Spikes in waterborne N. salmincola concentrations in October and November typically coincided with increases in streamflow; this combination resulted in elevated infection pressures during high water events in the fall. The peak in waterborne N. salmincola concentrations corresponded with an accompanying peak in tissue parasite density (metacercariae/posterior kidney) in Coho Salmon Oncorhynchus kisutch that were reared in the untreated water., (© 2019 American Fisheries Society.)

Digeneans (Trematoda: Digenea) are endoparasites that present a complex life cycle, generally involving an intermediate invertebrate host and a vertebrate host. There is limited information about which species of molluscs may act as intermediate hosts in the upper Paraná River floodplain (UPRF), where Aylacostoma chloroticum can be considered a potential candidate. The study of digeneans in this region is important because some of these parasites are potentially zoonotic, and, therefore, are relevant to public health. However, the correct identification of these organisms during the larval stages is difficult because of the lack of morphologically diagnostic characteristics. The objective of this study was to identify and molecularly characterize the larval stages of digeneans found in A. chloroticum in the UPRF, using the mitochondrial marker of subunit I of cytochrome c oxidase and the 28S nuclear marker. The molluscs were examined in the laboratory and three morphotypes of cercariae were found. DNA was extracted from the specimens obtained and was then amplified and sequenced. The morphotypes exhibited high genetic similarities with Pseudosellacotyla, Paralecithodendrium and Philophthalmus, indicating that these organisms belong to these genera. This is the first record of larval stages of these genera in molluscs collected in the UPRF.

Urban agriculture is very common in the cities, as Rio de Janeiro and adjacencies, being important as economic activity. However, this practice can create appropriate conditions for establishment and maintenance of life cycle of many parasites. Lymneid snails can act as intermediate hosts of many trematodes species, including those that infect humans and wild and domestic. In the present study, the snail Lymnaea columella was collected from an urban watercress plantations and the morphology of cercariae released by them was analyzed by light and scanning electron microscopy. The specimens were identified as xiphidiocercariae, belonging to the Cercariae Ornatae group, of the order Plagiorchiiformes. Histological examination of the digestive gland of the infected snails revealed the presence of developing sporocysts, with hemocitary infiltration and metaplasia in the epithelial tissue.

Objectives: To optimise host-to-host transmission, digenean trematodes (parasites) synchronise their cercarial emission patterns with the aquatic activities of their vertebrate hosts. Schistosoma mansoni has two different circadian chronotypes: a diurnal shedding pattern with a mean peak occurring at 11:00 h, and a nocturnal shedding pattern with a mean peak occurring at 20:00 h. We analysed the life-history variations between these two chronotypes at the levels of the parasite and its hosts., Methods: For each chronotype, we quantified three life-history traits related to the parasite (prepatent period, infection rate and cercarial production) and analysed the morphometry and the morphology of the parasite eggs; we also quantified three life-history traits related to the snail intermediate host (shell diameter, fecundity and survival rate) and one life-history trait related to the experimental definitive host (survival rate). A phylogeny based on the mitochondrial cytochrome-oxidase gene was made on samples of both chronotypes., Results: Life-history analysis revealed significant variations between the two chronotypes. Life-history traits were optimal for both the parasite and the snail host for the diurnal chronotype compared to the nocturnal one. The new chronotype behaved like an allopatric population towards its snail host. Phylogenetic analysis supports the hypothesis of a lateral transfer of S. mansoni from humans to Rattus rattus. These results were interpreted in terms of an ongoing sympatric speciation., Conclusion: The nocturnal chronotype of S. mansoni showed non-adapted life-history traits in its relation with the snail intermediate host Biomphalaria pfeifferi. The emergence of this new phenotype is probably linked to divergent natural selection., (© 2019 John Wiley & Sons Ltd.)

In the Upper Mississippi River Region, invasive faucet snails (Bithynia tentaculata) and their trematode parasites have been implicated in more than 182 000 waterfowl deaths since 1996. Estimating transmission potential depends on accurate assessments of susceptible host population size. However, little is known about the mechanisms underlying snail-host susceptibility in this system. Prior field studies suggest that very small, likely young, faucet snails are less suitable secondary intermediate hosts. Here, we test whether the patterns observed in the field are because small snails (1) are refractory to infection by cercariae, (2) die from infection and are removed from sampled populations, and/or (3) are not preferred by cercariae. Our own field collections were consistent with the observation that smaller faucet snails exhibit lower metacercarial infection prevalence and abundance than larger snails. However, laboratory-based experiments show that smaller snails were actually more susceptible to infection than larger snails. Moreover, the smallest snail size class had significantly higher mortality than larger snails following infection, which may explain their reduced infection levels observed in the field. Our study demonstrates the importance of pairing field and laboratory studies to better understand mechanisms underlying patterns of infection.

The potential for local biodiversity to affect transmission success of parasites has been shown to be particularly important in trematodes, where non-host organisms can feed on and 'dilute' free-living infective stages (cercariae). Earlier studies have analyzed the effects of various predators on transmission stages of single trematode species, but not how cercariae of different species react to predation pressure. Here, we tested whether cercariae with different host-searching movement patterns show varying susceptibility to predation by non-host species with different feeding habits. For this study, we performed a set of predation experiments with 6 species of trematode cercariae ( Coitocaecum parvum , Maritrema poulini , Apatemon sp., Telogaster opisthorchis , Plagiorchioid sp. I, and Aporocotylid sp. II) that represent 2 groups of host-searching behavior, free-swimming vs. bottom-dwelling, and 2 predators ( Sphaerium sp., Physa acuta ) with distinct feeding modes, a filter feeder and a grazer. Our results show that cercarial susceptibility to predation is highly dependent on the interspecific interaction between dispersal behavior of cercariae and feeding behavior of non-host organisms: Filter feeders only diluted free-swimming cercarial stages, not bottom-dwelling ones; grazers on the other hand, had no effect on free-swimming cercariae but reduced bottom-dwelling cercariae in 1 trematode species. Our findings give further support to the hypothesis that the transmission dynamics of trematodes do not simply depend on local biodiversity but rather on the species-specific interactions between parasite transmission stages and free-living organisms in the ecosystem. This has important implications for disease dynamics in ecological communities (e.g., the parasites' infection success), and for ecosystem energetics, as cercariae constitute important food items.

The ability of formalin, PEROX-AID (hydrogen peroxide), and seawater to kill waterborne Nanophyetus salmincola cercariae was evaluated in vitro. Newly emerged cercariae survived for extended periods in freshwater, with 53-73% survival occurring in negative control groups after 24 h. Exposure to dilutions of formalin reduced this survival time, with 0% of cercariae surviving after 30 min in 450 μL/L, 40 min in 225 μL/L, and 300 min in 113 μL/L. Exposure to PEROX-AID (hydrogen peroxide) for 1 h resulted in reduced cercarial survival (16.4%) only at the highest concentration (100 μL/L), compared with 100% survival in the untreated controls and all lesser concentrations. Exposure to dilutions of seawater resulted in reduced cercarial survival only at high salinities (15.2-30.3‰), where 10-min exposures resulted in 0-20% survival. These results provide insights into options for prophylactic water treatment at salmonid enhancement facilities that experience high mortalities due to infections with Nanophyetus salmincola. Further, the intolerance of live cercariae to high salinities indicates that exposure to fish occurs primarily in the freshwater portions of watersheds., (Published 2018. This article is a U.S. Government work and is in the public domain in the USA.)

A rich fauna of digenetic trematodes has been documented from the Great Barrier Reef (GBR), yet little is known of the complex life-cycles of these parasites which occur in this diverse marine ecosystem. At Heron Island, a small coral cay at the southern end of the GBR, the intertidal marine gastropod Clypeomorus batillariaeformis Habe & Kusage (Cerithiidae) is especially abundant. This gastropod serves as an intermediate host for 12 trematode species utilising both fish and avian definitive hosts. However, 11 of these species have been characterised solely with morphological data. Between 2015 and 2018 we collected 4870C. batillariaeformis from Heron Island to recollect these species with the goal of using molecular data to resolve their phylogenetic placement. We found eight of the 12 previously known species and two new forms, bringing the total number of digenean species known to parasitise C. batillariaeformis to 14. The families of this trematode community now include the Atractotrematidae Yamaguti, 1939, Bivesiculidae Yamaguti, 1934, Cyathocotylidae Mühling, 1898, Hemiuridae Looss, 1899, Heterophyidae Leiper, 1909, Himasthlidae Odhner, 1910, Microphallidae Ward, 1901, and Renicolidae Dollfus, 1939. Molecular data (ITS and 28S rDNA) were generated for all trematode species, and the phylogenetic position of each species was determined. The digenean community parasitising C. batillariaeformis includes several common species, as well as multiple species which are uncommon to rare. Although most of those trematodes in the community which exploit fishes as definitive hosts have remained common, the composition of those which utilise birds appears to have shifted over time., (Copyright © 2018 Elsevier B.V. All rights reserved.)

The transmission from one host to another constitutes a challenging obstacle for parasites and is a key determinant of their fitness. Due to their complex life histories involving several different hosts, the free-living dispersal stages (cercariae) of digenean trematodes show a huge diversity in morphology and behaviour. On a finer scale, we still have an extremely limited understanding of the inter- and intraspecific variation in transmission strategies of many trematode species. Here, we present a novel method to study the movement patterns of cercariae of four New Zealand trematode species (Coitocaecum parvum, Maritrema poulini, Apatemon sp. and Aporocotylid sp. I.) via automated video tracking. This approach allows to quantify parameters otherwise not measurable and clearly illustrates the individual strategies of parasites to search for their respective target hosts. Cercariae that seek out an evasive fish target hosts showed higher swimming speeds (acceleration and velocity) and travelled further distances, compared with species searching for high-density crustacean hosts. Automated video tracking provides a powerful tool for such detailed analyses of parasites' host-searching strategies and can enhance our understanding of complex host-parasite interactions, ranging from parasite community structure to the transmission of potential disease agents.

Background: Temperature is an important factor that influences the biology and ecology of intermediate host (IH) snails and the schistosome parasites they transmit. Although temperature shifts due to climate change has been predicted to affect the life history traits of IH snails and parasite production, the mechanisms of how this may affect parasite abundance and disease risks are still not clear., Materials and Methods: Using data from laboratory and field experiments, we developed a deterministic compartmental simulation model based on difference equations using a weekly time step that represented the life cycle of Bulinus globosus. We simulated snail population dynamics and the associated production of cercariae assuming current environmental temperatures as well as projected temperature increases of 1 °C and 2 °C., Results: The model generated snail fecundity and survival rates similar to those observed in the laboratory and also produced reasonable snail population dynamics under seasonally varying temperatures representative of generally favorable environmental conditions. Simulated relative abundances of both snails and cercariae decreased with increasing environmental temperatures, with maximum snail abundances decreased by 14% and 27%, and maximum cercariae productions decreased by 8% and 17%, when temperatures were increased by 1 °C and 2 °C, respectively., Conclusion: The results indicate that future rise in temperature due to climate change may alter the abundance of B. globosus and impact on the prevalence of schistosomiasis. Furthermore, increased temperatures may not linearly influence the abundance of S. haematobium. These results may have important implications for schistosomiasis control programmes in view of temperature driven changes in the life history traits of B. globosus and S. haematobium. Our study recommends that the use of deterministic models incorporating the effects of temperature on the life history traits of IH snails would be vital in understanding the potential impact of climate change on schistosomiasis incidences and prevalence., Competing Interests: The authors have declared that no competing interests exist.

The human liver fluke Opisthorchis viverrini (Platyhelminthes, Trematoda, Digenea) uses snails of the genus Bithynia as first intermediate host. Peculiarly among trematodes, the eggs of O. viverrini hatch within the digestive tract of its snail host. It remains uncertain whether hatching in this species is mediated through mechanical fracture of the eggshell or by digestion with specific digestive enzymes. This study aimed to characterize enzymes with specific inhibitors and factors involved in the hatching activity of O. viverrini eggs. For measuring egg hatching in vivo, 50 O. viverrini mature eggs were fed to individual Bithynia siamensis goniomphalos snails at various temperature conditions for 24 hr. Ex vivo, mature eggs were incubated with crude snail extract and commercial leucine aminopeptidase (LAP). Egg-hatching of O. viverrini was temperature dependent, with optimal hatching occurring at 24-28 C, with a peak of hatching of 93.54% in vivo and 30.55% ex vivo occurring at these temperatures. Ex vivo hatching rates increased to 45.87% under anaerobic conditions at 28 C. Some 22.70% and 16.21% of heat-killed eggs also hatched within the snail digestive tract and snail extract, respectively, indicating that host molecules are involved in the hatching response. Most eggs hatch in the anterior regions of the digestive tract. Hatching was completely inhibited in the presence of bestatin, an inhibitor of LAP, but not in the presence of phosphatase inhibitors. Bestatin inhibition of hatching was reversible. Finally, egg hatching could be induced by addition of a porcine LAP. The results indicate that this digenean utilizes both LAP of the snail host and movement of miracidia for hatching.

Parasites often use external cues to identify and move toward environments where they are likely to encounter suitable hosts. The trematode parasite Euhaplorchis californiensis produces cercariae that emerge from California horn snails ( Cerithideopsis californica [= Cerithidea californica]) to infect California killifish ( Fundulus parvipinnis) as second intermediate hosts. Based upon work on a congeneric Euhaplorchis species from Florida, and based on the ecology of its killifish host, we hypothesized that E. californiensis cercariae in southern California estuaries are positively phototactic and negatively geotactic, using both sunlight and gravity to guide their movement to the upper water column. To distinguish positive phototaxis from negative geotaxis, we first quantified E. californiensis movement in response to light along a horizontal plane and determined they were positively phototactic. In a second experiment, we quantified E. californiensis movement along a vertical plane in response to an overhead light, a light from below, or no light. We found that E. californiensis exhibit negative geotaxis in the absence of light, but will swim in the direction of gravity to move toward a light source from below. Thus, E. californiensis are both positively phototactic and negatively geotactic, but cercariae prioritize phototactic cues. These results suggest that E. californiensis cercariae aggregate in the open water, indicating that the pelagic zone represents an area of high infection risk for California killifish hosts.

Background: Schistosomiasis is one of the most disabling neglected tropical diseases, ranking second in terms of years lived with disability. While treatment with the drug praziquantel can have immediate beneficial effects, reinfection can occur rapidly if people are in contact with cercaria-infested water. Water treatment for schistosomiasis control seeks to eliminate viable cercariae from water, thereby providing safe alternative water supplies for recreational and domestic activities including laundry and bathing. This provision may reduce contact with infested water, which is crucial for reducing reinfection following chemotherapy and cutting schistosome transmission., Methodology: A qualitative systematic review was carried out to summarize the existing knowledge on the effectiveness of water treatment in removing or inactivating human schistosome cercariae. Four online databases were searched. Studies were screened and categorized into five water treatment processes: storage, heating, chlorination, filtration, and ultraviolet (UV) disinfection., Conclusions: All five water treatment methods can remove or inactivate cercariae in water, and hence produce cercaria-free water. However, reliable design guidelines for treating water do not exist as there are insufficient data. Overall, the review found that cercariae are inactivated when storing water for 10-72 hours (depending on temperature), or with chlorination values of 3-30 mg-min/l. UV fluences between 3-60 mJ/cm2 may significantly damage or kill cercariae, and sand filters with 0.18-0.35 mm grain size have been shown to remove cercariae. This systematic review identified 67 studies about water treatment and schistosomiasis published in the past 106 years. It highlights the many factors that influence the results of water treatment experiments, which include different water quality conditions and methods for measuring key parameters. Variation in these factors limit comparability, and therefore currently available information is insufficient for providing complete water treatment design recommendations.

Lecithodendrium linstowi is one of the most prevalent and abundant trematodes of bats, but the larval stages and intermediate hosts have not been identified. We present the first molecular and morphological characterization of the cercariae of L. linstowi based on a phylogenetic analysis of partial fragments of LSU and ITS2 rDNA. The first intermediate host was incriminated as Radix balthica by DNA barcoding using cox1 and ITS2 sequences, although the snail morphologically resembled Radix peregra, emphasizing the requirement for molecular identification of lymnaeids as important intermediate hosts of medical and veterinary impact. The application of molecular data in this study has enabled linkage of life cycle stages and accurate incrimination of the first intermediate host.

Recent results suggest that bivalves can play an important role in restraining the spread of various aquatic infections. However, the ability of mussels to remove free-living stages of macroparasites and reduce their transmission is still understudied, especially for freshwater ecosystems. We investigated the influence of the common freshwater mussel (Anodonta anatina) on the transmission of a trematode (eye fluke, Diplostomum pseudospathaceum), which frequently infects fish in farms and natural habitats. In our experiments, mussels caused a significant decrease (P < 0·001) in the abundance of trematode free-living stages, from 6520 to 1770 cercariae L-1 on average (about 4-fold in 2 h). Individual clearance rates of mussels were 0·6‒3·7 L per hour (mean 1·9). These tests were followed by experimental infections of rainbow trout (Oncorhynchus mykiss) with different doses of D. pseudospathaceum cercariae in the presence or absence of mussels. Exposure of fish to cercariae in the presence of mussels significantly (P < 0·05) reduced the infection intensities in fish (by 30-40%) at all exposure doses. Our results indicate that freshwater bivalves can markedly reduce local cercariae densities and could be useful in mitigation of trematodoses harmful to fish farming.

In parasites with complex life cycles the transmission of free-living infective stages can be influenced by ambient community diversity, in particular via predation. Here, we experimentally investigated whether parasite density and the presence of alternative prey can alter predation rates on free-living cercarial stages of a marine trematode by several non-host predators. All four predator species consumed increasing numbers of cercariae with an increase in cercarial density, indicating that the removal of cercariae by predators is effective over a range of natural densities as well as in the presence of alternative prey for a number of predators typical of marine ecosystems. However, the relative removal rates and the effects of cercarial density and alternative prey differed among predator species. In barnacles and shrimps, significant interactive effects of cercarial density and alternative prey on cercarial predation occurred while in oysters and crabs cercarial removal rates were unaffected by both factors. As changes in cercarial densities directly translate into changes in infection levels in down-stream hosts in this parasite-host system, the observed predator-specific responses suggest that cercarial predation effects on disease risks will depend on the specific species composition of ambient communities and not on non-host biodiversity per se.