Description: While the free-living fauna of the Wadden Sea has received much interest, little is known on the distribution and effects of parasites in the Wadden Sea food web. However, recent studies on this special type of trophic interaction indicate a high diversity of parasites in the Wadden Sea and suggest a multitude of effects on the hosts. This also includes effects on specific predator-prey relationships and the general structure of the food web. Focussing on molluscs, a major group in the Wadden Sea in terms of biomass and abundance and an important link between primary producers and predators, we review existing studies and exemplify the ecological role of parasites in the Wadden Sea food web. First, we give a brief inventory of parasites occurring in the Wadden Sea, ranging from microparasites (e.g. protozoa, bacteria) to macroparasites (e.g. helminths, parasitic copepods) and discuss the effects of spatial scale on heterogeneities in infection levels. We then demonstrate how parasites can affect host population dynamics by acting as a strong mortality factor, causing mollusc mass mortalities. In addition, we will exemplify how parasites can mediate the interaction strength of predator-prey relationships and affect the topological structure of the Wadden Sea food web as a whole. Finally, we highlight some ongoing changes regarding parasitism in the Wadden Sea in the course of global change (e.g. species introduction, climate change) and identify important future research questions to entangle the role of parasites in the Wadden Sea food web

Description: In the Western Baltic Sea, climate change is happening at much faster rate than in most other seas and organisms are additionally exposed to a steep and variable salinity gradient. Climate change has previously been shown to affect parasite transmission in other marine ecosystems, yet little is known about potential effects of warming and desalination on parasite–host interactions. In laboratory experiments, we determined the combined effects of projected seawater warming and freshening on the emergence, activity, survival, and infectivity of cercariae (free-swimming infectious stage) of the trematode Himasthla elongata (Mehlis 1831), shed from its first intermediate host, the periwinkle Littorina littorea (Linnaeus 1758), in the Baltic Sea. We also assessed the susceptibility of the second intermediate host, the mussel Mytilus edulis Linnaeus, 1758, to cercarial infections. Generally, salinity was the main driver, particularly of cercarial activity, infectivity, and mussel susceptibility to infection. At the lowest salinity (13), cercariae were 50% less active compared to the highest salinity (19). Infection success and host susceptibility followed a similar pattern, with 47% and 43% less metacercariae (encysted stage) present at salinity 13 than at salinity 19, respectively. In contrast, effects of simulated warming were found only for cercarial survival, with cercarial longevity being higher at 19 than at 23 °C. No significant interactions between temperature and salinity were found. In contrast to the literature, the results suggest that a climate change-driven freshening (partly also warming) may lead to a general decline of marine trematodes, with possible beneficial effects for the involved hosts.

Description: Trematode prevalence and abundance in hosts are known to be affected by biotic drivers as well as by abiotic drivers. In this study, we used the unique salinity gradient found in the south-western Baltic Sea to: (i) investigate patterns of trematode infections in the first intermediate host, the periwinkle Littorina littorea and in the downstream host, the mussel Mytilus edulis, along a regional salinity gradient (from 13 to 22) and (ii) evaluate the effects of first intermediate host (periwinkle) density, host size and salinity on trematode infections in mussels. Two species dominated the trematode community, Renicola roscovita and Himasthla elongata. Salinity, mussel size and density of infected periwinkles were significantly correlated with R. roscovita, and salinity and density correlated with H. elongata abundance. These results suggest that salinity, first intermediate host density and host size play an important role in determining infection levels in mussels, with salinity being the main major driver. Under expected global change scenarios, the predicted freshening of the Baltic Sea might lead to reduced trematode transmission, which may be further enhanced by a potential decrease in periwinkle density and mussel size.

Description: Global warming may alter the dynamics of infectious diseases by affecting important steps in the transmission of pathogens and parasites. In trematode parasites, the emergence of cercarial stages from their hosts is temperature-dependent, being highest around a thermal optimum. If environmental temperatures exceed this optimum as a consequence of global warming, this may affect cercarial transmission. However, our knowledge of cercarial emergence patterns of species from high temperature environments is currently very limited. Here, we investigated the effect of temperature on the emergence of two common trematode species from an abundant mud snail Pirenella cingulata in the Persian Gulf, the warmest sea on Earth. Infected snails were incubated in the laboratory at 6 temperatures from 10 to 40°C for 3 days. We found an optimal temperature for cercarial emergence of 32.0°C and 33.5°C for Acanthotrema tridactyla and Cyathocotylidae gen. sp., respectively, which are the warmest recorded thermal optima for any aquatic trematode species. Emergence of both species dropped at 40°C, suggesting upper thermal limits to emergence. Overall, Persian Gulf trematodes may be among the most heat-tolerant marine trematode species, indicating a potential for dispersing to regions that will continue to warm in the future.

Description: Predators can affect parasite–host interactions when directly preying on hosts or their parasites. However, predators may also have non-consumptive indirect effects on parasite–host interactions when hosts adjust their behaviour or physiology in response to predator presence. In this study, we examined how chemical cues from a predatory marine crab affect the transmission of a parasitic trematode from its first (periwinkle) to its second (mussel) intermediate host. Laboratory experiments revealed that chemical cues from crabs lead to a threefold increase in the release of trematode cercariae from periwinkles as a result of increased periwinkle activity. This positive effect on transmission was contrasted by a 10-fold reduction in cercarial infection rates in the second intermediate host when we experimentally exposed mussels to cercariae and predator cues. The low infection rates were caused by a substantial reduction in mussel filtration activity in the presence of predator cues, preventing cercariae from entering the mussels. To assess the combined net effect of both processes, we conducted a transmission experiment between infected periwinkles and uninfected mussels. Infection levels of mussels in the treatments with crab cues were sevenfold lower than in mussels without crab chemical cues. This suggests that predation risk effects on mussel susceptibility can counteract the elevated parasite release from first intermediate hosts, with negative net effects on parasite transmission. These experiments highlight that predation risk effects on parasite transmission can have opposing directions at different stages of the parasite's life cycle. Such complex non-consumptive predation risk effects on parasite transmission may constitute an important indirect mechanism affecting prevalence and distribution patterns of parasites in different hosts across their life cycle.

Description: The mussel Mytilus edulis, a host to various trematode species, experiences performance decrements due to these infections. Yet, the impact magnitude and potential interactions with environmental stressors remain largely unexplored. This study scrutinizes the effect of Renicola roscovita infections on mussel filtration and respiration. We first assessed performance in both uninfected and lab-infected mussels at a mild temperature (16 °C), following an acute heat ramp to 30.5 °C and subsequent cooling. The experiment revealed neither a significant direct impact of the infection on the mussels' performance, nor any significant interplay between the infection and temperature variations. To account for possible infection effects obscured by low sample sizes or mussel size disparities, we conducted a reassessment at 16 °C using both small and large mussels. Infection notably hampered filtration in large mussels, with a marginal impact on smaller ones. A positive correlation was found between infection intensity and mussel filtration capacity, though the infection had no discernible impact on respiration. Our consistent finding of an 11-12 % infection effect size across all experiments indicates a slight reduction in mussel filtration due to trematode infections. While the exacerbating effect of transient heat stress on the infection's impact on filtration was not statistically significant, future investigations should explore potential interactions with prolonged heat stress. Our findings underscore the nuanced ways in which parasitic infections can influence marine bivalve physiology, emphasizing the need for more comprehensive studies that incorporate environmental stressors, such as heat stress, to fully elucidate the impact of parasitism on marine ecosystem health and resilience.

Description: Avian schistosomes, comprise a diverse and widespread group of trematodes known for their surprising ability to switch into new hosts and habitats. Despite the considerable research attention on avian schistosomes as causatives of the human cercarial dermatitis, less it is known about the diversity, geographical range and host associations of the marine representatives. Our molecular analyses inferred from cox1 and 28S DNA sequence data revealed presence of two schistosome species, Ornithobilharzia canaliculata (Rudolphi, 1819) Odhner, 1912 and a putative new species of Austrobilharzia Johnston, 1917. Molecular elucidation of the life-cycle of O. canaliculata was achieved for the first time via matching novel and published sequence data from adult and larval stages. This is the first record of Ornithobilharzia from the Persian Gulf and globally the first record of this genus in a potamidid snail host. Our study provides: (i) new host and distribution records for major etiological agents of cercarial dermatitis and contributes important information on host-parasite relationships; (ii) highlights the importance of the molecular systematics in the assessment of schistosome diversity; and (iii) calls for further surveys to reach a better understanding of the schistosome diversity and patterns of relationships among them, host associations, transmission strategies and distribution coverage.

Description: In this experiment, we assessed the different transmission steps from the first to the second intermediate host: i) cercarial emergence from periwinkles, ii) cercarial activity and survival after emergence, iii) cercarial infectivity in mussels, and iv) susceptibility of mussels to cercarial infection. For iii) cercariae were treated but not the mussels, whereas for iv) mussels were treated but not the cercariae. The experiment was run in August-September 2017 in the climate chambers of GEOMAR in Kiel. All experiments were conducted using temperature and salinity (fully crossed) as well as time (only for cercarial output) as fixed factors, and periwinkle/mussel identity nested within water bath as random factor. Temperature levels applied were 19 and 23°C. Salinity levels applied were 13, 16 and 19.

Description: Cercarial emergence was assessed by counting all cercariae that emerged from single periwinkles exposed to the different treatments. Periwinkles were collected in Årøsund, Denmark and screened in the laboratory to verify the infection status. To measure cercarial emergence, periwinkles were individually transferred from the jars into 50 mL Plexiglas beakers, one day after the 7-days acclimation (sampling 1), one week after the start of exposure to the treatments (sampling 2), and two weeks after the start of exposure to the treatments (sampling 3). Each incubation lasted eight hours. At the end of the incubation period, the water from each beaker was transferred into 50 mL Falcon tubes. In addition, the beakers were immediately washed with 5 mL ethanol (99%), and the solution was also added to the Falcon tube to ensure the collection of all cercariae and their preservation for later counting.

Description: For cercarial activity and survival infected periwinkles were acclimated for three weeks under the different treatments (n=10 individuals per treatment) and incubated to allow for cercarial release. At the start of the experiment, approximately 45 cercariae per treatment were added individually to wells of three 96-well plates The 96-well plates were then exposed to the different temperature treatments placing them in the six thermobaths (three 96-well plates per treatment combination). Activity and survival were then assessed under a stereomicroscope after 4, 6, 9, 18, 27, and 45 hours. Cercariae were considered fully active when they were constantly swirling around (category: fully active). When laying at the bottom and not reacting, these were considered dead (category: dead), when still reacting, even if slowly, these were still considered alive (category: alive). This last category included "fully active".