Data on distribution, demographic structure and grazing of the dominant mesozooplankton species in the Yenisei estuary and adjacent shelf in early summer.

The data article refers to the paper "Distribution and grazing of the dominant mesozooplankton species in the Yenisei estuary and adjacent shelf in early summer (July 2016)" (Drits et al., 2020). The data were collected along quasi-longitudinal transect "Yenisei estuary - Kara Sea shelf" on 24-28 July 2016. Here we present data on the spatial and vertical distribution, demographic structure and gut pigment content of the dominant zooplankton species as well as the grazing impact on autotrophic phytoplankton in the three distinguished zones: freshwater zone, frontal zone of the Yenisei plume and marine shelf zone. The related article (Drits et al., 2020) considers the structure and functioning of zooplankton community in relation to environmental characteristics such as temperature, salinity, phytoplankton abundance, timing of ice retreat. Information presented in this article can be used by marine biologists for studies of structure and functioning of estuarine pelagic communities, ecology of zooplankton in the Siberian seas. Besides the data could provide a baseline for the assessment of the ecological role played by climate change events (e.g., increased precipitation, permafrost thawing, elevated river discharge) on the Arctic ecosystems.

Personality Influences Risk of Parasitism in Fish.

Influence of fish personality on infection rate is poorly studied. In the experiments on young-of-the-year Oncorhynchus mykiss and cercariae of the trematode Diplostomum pseudospathaceum, we tested the hypothesis that infection rate differs between more and less active ("bold" and "shy") fish. Will individual differences in infection persist upon re-infection? Fish serve as a second intermediate host for this trematode. Positive correlation was found between the results of consecutive infections. Accumulation of parasites with successive infections leads to an aggregated distribution of D. pseudospathaceum among the hosts, affecting individual fitness and polymorphism in fish populations. Persistent individual differences in parasite burden among fish and, as a result, vulnerability for predators confirms the role of parasites as an important factor of natural selection.

Zooplankton impact on the organic matter flux of Siberian Arctic seas.

Role of zooplankton in the organic matter flux was studied in the Kara and Laptev seas using sediment traps in the course of voyage 63 of the Academician Mstislav Keldysh research vessel in August-October 2015. The values of the total flux and that of organic matter were at least an order of magnitude higher than those obtained before the 2000s. A hypothesis is proposed on an increasing influence of the river runoff on the Kara Sea area under climatic changes. Zooplankton contribution into Corg flux varied broadly, averaging 50%. The highest flux values and contribution of zooplankton in them (up to 96%) were recorded in the frontal (gradient) zones.

Behavioural adaptations of argulid parasites (Crustacea: Branchiura) to major challenges in their life cycle.

Fish lice (Argulus spp.) are obligate ectoparasites, which contrary to most aquatic parasites, retain the ability to swim freely throughout the whole of their life. In fish farms, they can quickly increase in numbers and without effective control cause argulosis, which results in the reduced growth and survival of their fish hosts. The morphology of Argulus spp, including their sensory organs, is suitable for both parasitism and free-swimming. By spending a considerable amount of time away from their host, these parasites risk being excessively dispersed, which could endanger mating success. Here we present a review of recent studies on the behaviour of Argulus spp, especially the aggregative behaviour that mitigates the dilution of the parasite population. Aggregation of parasites, which is especially important during the period of reproduction, occurs on different scales and involves both the aggregation of the host and the aggregation of the parasites on the host. The main behavioural adaptations of Argulus spp, including searches for hosts and mates, host manipulation and host choice, are all focused on the fish. As these ectoparasites repeatedly change hosts and inflict skin damage, they can act as vectors for fish pathogens. The development of environmentally friendly measures for the control and prevention of argulosis needs to take into account the behaviour of the parasites.

Parasite-induced aggression and impaired contest ability in a fish host.

BACKGROUND: Success of trophically transmitted parasites depends to a great extent on their ability to manipulate their intermediate hosts in a way that makes them easier prey for target hosts. Parasite-induced behavioural changes are the most spectacular and diverse examples of manipulation. Most of the studies have been focused on individual behaviour of hosts including fish. We suggest that agonistic interactions and territoriality in fish hosts may affect their vulnerability to predators and thus the transmission efficiency of trophically transmitted parasites. The parasite Diplostomum spathaceum (Trematoda) and juvenile rainbow trout, Oncorhynchus mykiss, were used to study whether infection can alter aggression rates and territorial behaviour of intermediate fish hosts. RESULTS: The changes in behaviour of rainbow trout, Oncorhynchus mykiss, infected with an eye fluke Diplostomum spathaceum (Trematoda), was monitored over the course of an experimental infection for 1.5 months. At the beginning of their development, not yet infective D. spathaceum metacercariae decreased the aggressiveness of rainbow trout. By the time that metacercariae were fully infective to their definitive hosts, the aggressiveness increased and exceeded that of control fish. Despite the increased aggressiveness, the experimentally infected fish lost contests for a territory (dark parts of the bottom) against the control fish. CONCLUSIONS: The results obtained indicate that the parasitized fish pay the cost of aggressiveness without the benefit of acquiring a territory that would provide them with better protection against predators. This behaviour should increase transmission of the parasite as expected by the parasite manipulation hypothesis.

Host specificity of Argulus coregoni (Crustacea: Branchiura) increases at maturation.

We tested the hypothesis that host specificity in ectoparasites does not depend exclusively on the features of the host but also on surrounding habitats, using 2 fish ectoparasites, Argulus coregoni and A. foliaceus (Crustacea: Branchiura), occurring sympatrically in Finnish lakes. Although these parasites are considered to be of low specificity, we found that the larger of the 2 species, A. coregoni developed a pronounced preference for salmonid hosts at the beginning of maturation (defined by the presence of copulating specimens). Argulus foliaceus infects a much wider range of fish hosts. We showed that specialization of A. coregoni on salmonids does not necessarily result from incompatibility with other fishes, but could instead reflect higher sensitivity of oxygen depletion compared with A. foliaceus. Adult A. coregoni may meet these demands by attaching to salmonids, the typical inhabitants of well-aerated waters. Young parasites of both species showed little host specificity and attached mainly to fishes with higher body reflectivity. In host choice experiments, A. coregoni of 4-5 mm length preferred salmonids (rainbow trout) to cyprinids (roach) irrespective of the type of fish host, on which it had been previously grown in the laboratory. We suggest that such an innate ontogenetic shift in host preference maintains the major part of the parasite population on its principal host, ensuring successful reproduction within suitable habitats.

Seasonal dynamics of egg laying and egg-laying strategy of the ectoparasite Argulus coregoni (Crustacea: Branchiura).

Substrate preferences, spatial aggregation patterns and seasonal dynamics in the egg laying of ectoparasitic Argulus coregoni were studied at a commercial fish farm in Finland. Pilot experiments showed that A. coregoni females selected specific types of substrates for egg laying. Significantly more A. coregoni eggs were laid on dark substrates than on light ones suggesting the use of visual cues. Therefore, egg-laying plates of dark colour were constructed for further experiments. Most A. coregoni eggs were deposited in locations in shadow and in the deepest water in a 2 m deep farming canal. Relatively more eggs were laid on bottom stones situated near each egg-laying trap than on artificial egg-laying plates indicating a preference for irregular stones in the deeper locations in the canal. The plates were located 20 cm above the bottom. However, a total of 5,863 A. coregoni egg clutches, corresponding approximately to 1.5 million unhatched metanauplii, were successfully destroyed with the plates indicating that egg-laying traps can be used as an ecological control method against argulids in certain situations. For traps to be effective, ponds should not contain stones or any other hard substrata attracting female lice. The egg laying of A. coregoni in this study started on 5 July in 2001 and extended over 3.5 months up to mid-October. The egg-laying pattern of A. coregoni population was unimodal, supporting the view that only a single A. coregoni generation occurred annually in Central Finland.

How do fish ectoparasites Argulus spp. (Crustacea: Branchiura) match with their hosts at the behavioural and ecological scales?

Features of the Argulus foliaceus and A. coregoni life cycles including spatial distribution and hatching patterns of the resting eggs as well as their host searching behaviour were analyzed on the basis of field and experimental data obtained in lakes and a fish farm in the Central Finland. Published data on behaviour and ecology of these parasites were also used. The main point of this analysis was an ecological adjustment of the life cycles of parasites to that of their fish hosts at different spatial and temporal scales. Due to different spatial distribution of overwintering eggs, the newly hatched larvae of A. coregoni are more likely to attach to salmonids, which are their main hosts, while A. foliaceus to percids and cyprinids. The main peak of argulid hatching occurs in late spring-early summer and coincides with the period when most of freshwater fishes concentrate close to the shore, either for spawning, or, foraging. Nevertheless, despite the general coincidence, the parasite-host encounter rate could still be very low. The main peak of hatching is followed by a long "tail". We suggest that under fluctuating and uncertain environmental conditions characterised by both abiotic stochasticity and poor predictability of the hosts, an optimal recruitment strategy is to hedge a bet, with a proportion of eggs synchronously hatching and a proportion remaining dormant and hatching at much lower rate during an extended period. One of the main behavioural adaptations to match with the hosts is all-over-the-day searching activity of argulids. The boreal A. coregoni relies primarily on vision, which is most effective in transparent waters during long polar days in summer. Vision is also important in the behavioural repertoire of the temperate A. foliaceus in the light, but olfaction and mechanoreception in the dark provide the highest host searching rate. The two generally sympatric species of argulids segregated on the finer habitat scale. Partially overlapping on salmonid hosts, they prefer ecologically distinct categories of fish, and use different microhabitats for egg laying and searching for hosts.

Spatial distribution and hatching of overwintered eggs of a fish ectoparasite, Argulus coregoni (Crustacea: Branchiura).

The habitat distribution of overwintered eggs, which were found to be the only source of spring recruitment of Argulus coregoni Thorell, was studied at a commercial fish farm in Central Finland. The frequency of occurrence of egg clutches in the deep parts of the canals and ponds was 50 to 80% and the percentage cover of the surface of stones with egg clutches was 1.7 to 6.4%, while in the shallow parts these values were 8 to 27% and 0.1 to 0.3%, respectively. A greater proportion of empty egg-shells was observed in shallow water in the mid-May, suggesting an earlier hatching there stimulated by the increased temperature and higher illumination. Under laboratory conditions, only elevated UV illumination, but not diurnally fluctuating temperature, significantly accelerated hatching. Normally overwintered eggs produced a pronounced peak of hatched larvae at the end of May and hatching continued at a much slower rate throughout the summer. Eggs that overwintered twice, first normally and then for a second time buried under sediments, were exposed to the same laboratory conditions simultaneously with normally overwintered eggs, but their hatching was delayed until August. The hatching rate was low, but markedly increased in December.

Effect of Triaenophorus crassus (Cestoda) infection on behavior and susceptibility to predation of the first intermediate host Cyclops strenuus (Copepoda).

Some parasites have been shown to manipulate host behavior so that parasite transmission to the next host is enhanced. Infection with Triaenophorus crassus Forel (Cestoda) caused alterations in the activity and microhabitat selection of the first intermediate host Cyclops strenuus Fischer (Copepoda) in the laboratory. Infected copepods made more starts to swim but spent less time swimming than uninfected copepods. These changes were independent of the intensity of infection. In a water column illuminated from above, infected copepods approached the surface, whereas uninfected ones remained close to the bottom. In the dark both infected and uninfected copepods stayed near the bottom. Finally, infection with T. crassus increased the probability of C. strenuus being eaten by the second intermediate host, whitefish (Coregonus lavaretus L. s.l.), in the laboratory. In experimental infections, 10-day-old procercoids had significantly lower infectivity for whitefish than older (12-, 14-, and 21-day-old) procercoids. Behavioral changes were detected in infected copepods containing procercoids 12 days old or older but not in experiments with 10-day-old procercoids. These results may indicate that T. crassus changes the behavior of the copepod host only after it has become infective to the next host, which is consistent with the active manipulation hypothesis.

Light-mediated host searching strategies in a fish ectoparasite, Argulus foliaceus L. (crustacea: branchiura).

Argulus foliaceus, an obligate fish ectoparasite, can search for its hosts in both light and dark conditions and uses vision in the light. We have examined what searching mode is used at night, when the infection rate was at its highest, and which stimuli produced by the fish are most important. A change of illumination produced a clear difference in the searching behaviour of adult Argulus females. The mean swimming speed and the area explored were 3-4 times higher in the dark, when the parasite employed a cruising search strategy. This changed to an ambush (hover-and-wait) strategy in the light. The swimming activity is accompanied by changes in metabolic costs; the activity of the electron transport system being approximately 25 % lower in the light. The most pronounced light-induced differences in host-searching behaviour took place in moderately hungry parasites (starved for 24-96 h). Less motivated (just having left a fish) or exhausted animals did not exhibit any clear differences in swimming speed. Among the external signals tested, fish smell, from both perch (Perca fluviatilis) and roach (Rutilus rutilus), induced an elevated swimming speed of the parasite. Periodic water movements caused similar but weaker effects. The effects of these stimuli were observed under both light and dark conditions. We conclude that host-searching behaviour of A. foliaceus is under internal (state of hunger) and external (illumination and host-induced signals) control and involves all its sensory equipment (vision, olfaction and mechano-reception). Perch (but not roach) reduced their swimming speed in the dark, which make them more susceptible to cruising Argulus. Thus the behavioural interplay between hosts and parasites can also influence the infection rate of A. foliaceus found on perch and roach in Finnish lakes.

Factors affecting abundance of Triaenophorus infection in Cyclops strenuus, and parasite-induced changes in host fitness.

Factors affecting the abundance of Triaenophorus crassus and Triaenophorus nodulosus procercoids in their copepod first intermediate host, Cyclops strenuus, and effects of infection on feeding behaviour, reproduction and survival of the host were studied experimentally. When exposed to the same number of coracidia, copepods harboured considerably less procercoids in the trials where ciliates or Artemia salina nauplii were given as alternative food items. The prevalence of infection was higher in adult copepods as compared with copepodite stage IV and stage V, and higher in stage V than in stage IV. The prevalences in adult females and males did not differ significantly from each other. The frequency of females carrying egg sacs was lower among infected than among exposed uninfected and unexposed copepods. The rate of feeding on Artemia nauplii remained at the same level in uninfected copepods, but decreased strongly in infected copepods during 7 days p.i. The survival of unexposed, exposed uninfected and infected copepods did not differ significantly from each other for the first 11 days post-exposure, but the mortality of infected copepods increased significantly after 3 weeks post-exposure. However, the rate of development and mortality of copepods might have been affected by the apparently arrested development of stage IV copepodites found in the experiment. Some of the contradictions between these results and earlier observations are suggested to be caused by the differences in the duration of exposure, intensity of infection and duration of observation post-exposure in the present study as compared with other experiments.

Changes in metabolism and behaviour of the freshwater copepod Cyclops strenuus abyssorum infected with Diphyllobothrium spp.

In a population of copepods (Cyclops strenuus abyssorum), with a naturally high prevalence of infection with procercoids of Diphyllobothrium spp., no difference in body size was found between infected and uninfected hosts. However, a significant reduction in the reproductive capacity of infected females was observed, 87% of uninfected females having developed eggs in their gonads and sacs compared with 21% of infected females. The feeding rate of infected copepods was relatively high soon after infection occurred, but gradually decreased to less than half that of uninfected animals. Respiration rate was also lower in infected copepods. Infected copepods showed reduced motility and impaired escape responses, which is likely to make them more susceptible to predation.