How cunning is the puppet-master? Cestode-infected fish appear generally fearless.

Trophically transmitted parasites have life cycles that require the infected host to be eaten by the correct type of predator. Such parasites should benefit from an ability to suppress the host's fear of predators, but if the manipulation is imprecise the consequence may be increased predation by non-hosts, to the detriment of the parasite. Three-spined sticklebacks (Gasterosteus aculeatus) infected by the cestode Schistocephalus solidus express reduced antipredator behaviours, but it is unknown whether this is an example of a highly precise manipulation, a more general manipulation, or if it can even be attributed to mere side effects of disease. In a series of experiments, we investigated several behaviours of infected and uninfected sticklebacks. As expected, they had weak responses to simulated predatory attacks compared to uninfected fish. However, our results suggest that the parasite induced a general fearlessness, rather than a precise manipulation aimed at the correct predators (birds). Infected fish had reduced responses also when attacked from the side and when exposed to odour from a fish predator, which is a "dead-end" for this parasite. We also tested whether the reduced anti-predator behaviours were mere symptoms of a decreased overall vigour, or due to parasite-induced hunger, but we found no support for these ideas. We propose that even imprecise manipulations of anti-predator behaviours may benefit parasites, for example, if other behaviours are altered in a way that increases the exposure to the correct predator.

Bitten or struck by dog: A rising number of fatalities in Europe, 1995-2016.

We analyzed fatal dog attacks in Europe 1995-2016 using official death cause data from Eurostat. The data comprised the number of fatalities assigned The International Statistical Classification of Diseases and Related Health Problems, Tenth Revision (ICD-10) code W54 "bitten or struck by dog", which includes deaths due to direct attacks but which excludes many complications following dog bites, such as rabies. In 2016, dogs killed 45 Europeans, which translates to an incidence of 0.009 per 100,000 inhabitants. This is comparable to estimates from the USA (0.011), and Canada (0.007), but higher than Australia (0.004). The number of European fatalities due to dog attacks increased significantly at a rate of several percent per year. This increase could not be explained by increases in the human or the dog populations. By taking all fatalities reported 1995-2016 into account, we investigated the effects of age, gender and geography. First, children, including infants, were common victims, but also middle-aged and the elderly, while people between ages 10 and 39 were rarely killed by dogs. Second, boys and men were overrepresented, but only in certain age groups and in certain parts of Europe. Third, there were large national and regional differences, both in the effects of gender and in incidences, which ranged from 0 to 0.045 per 100,000 inhabitants. This study of dog-related fatalities at a European level is the first of its kind and forms a basis for more detailed, national studies.

Phenotypic flexibility in background-mediated color change in sticklebacks.

Phenotypic flexibility may incur a selective advantage in changing and heterogeneous environments, and is increasingly recognized as an integral aspect of organismal adaptation. Despite the widespread occurrence and potential importance of rapid and reversible background-mediated color change for predator avoidance, knowledge gaps remain regarding its adaptive value, repeatability within individuals, phenotypic correlates, and whether its expression is context dependent. We used manipulative experiments to investigate these issues in two fish species, the three-spined stickleback (Gasterosteus aculeatus) and nine-spined stickleback (Pungitius pungitius). We sequentially exposed individuals to dark and light visual background treatments, quantified color change from video recordings, and examined associations of color change with phenotypic dimensions that can influence the outcome of predator-prey interactions. G. aculeatus expressed a greater degree of color change compared to P. pungitius. In G. aculeatus, the color change response was repeatable within individuals. Moreover, the color change response was independent of body size but affected by sex and boldness, with males and bolder individuals changing less. Infection by the parasite Schistocephalus solidus did not affect the degree of color change, but it did modulate its association with sex and boldness. G. aculeatus adjusted the expression of color change in response to predation risk, with enhanced color change expression in individuals exposed to either simulated attacks, or olfactory cues from a natural predator. These results provide novel evidence on repeatability, correlated traits, and context dependence in the color change response and highlight how a suite of factors can contribute to individual variation in phenotypic flexibility.

Aggressive desert goby males also court more, independent of the physiological demands of salinity.

Both between- and within-individual variation in behaviour can be important in determining mating opportunities and reproductive outcomes. Such behavioural variability can be induced by environmental conditions, especially if individuals vary in their tolerance levels or resource allocation patterns. We tested the effects of exposure to different salinity levels on male investment into two important components of mating success-intrasexual aggression and intersexual courtship-in a fish with a resource defence mating system, the desert goby, Chlamydogobius eremius. We found that males that were more aggressive to rivals also exhibited higher rates of courtship displays towards females. Contrary to predictions, this positive relationship, and the consistency of the two behaviours, were not affected by the salinity treatment, despite the physiological costs that high salinity imposes on the species. Moreover, over the entire data-set, there was only a marginally non-significant tendency for males to show higher levels of aggression and courtship in low, than high, salinity. The positive correlation between male aggression and courtship, independent of the physiological demands of the environment, suggests that males are not inclined to make contrasting resource investments into these two key reproductive behaviours. Instead, in this relatively euryhaline freshwater species, typical investment into current reproductive behaviours can occur under a range of different salinity conditions.

Consequences of eye fluke infection on anti-predator behaviours in invasive round gobies in Kalmar Sound.

Larvae of the eye fluke, Diplostomum, emerge from snails and infect fish by penetrating skin or gills, then move to the lens where they may impair the vision of the fish. For the fluke to reproduce, a bird must eat the infected fish, and it has been suggested that they therefore actively manipulate the fish's behaviour to increase the risk of predation. We found that round gobies Neogobius melanostomus, a species that was recently introduced to the Kalmar Sound of the Baltic Sea, had an eye fluke prevalence of 90-100%. We investigated how the infection related to behavioural variation in round gobies. Our results showed that the more intense the parasite-induced cataract, the weaker the host's response was to simulated avian attack. The eye flukes did not impair other potentially important anti-predator behaviours, such as shelter use, boldness and the preference for shade. Our results are in accordance with the suggestion that parasites induce changes in host behaviour that will facilitate transfer to their final host.

The influence of recent social experience and physical environment on courtship and male aggression.

BACKGROUND: Social and environmental factors can profoundly impact an individual's investment of resources into different components of reproduction. Such allocation trade-offs are expected to be amplified under challenging environmental conditions. To test these predictions, we used a desert-dwelling fish, the desert goby, Chlamydogobius eremius, to experimentally investigate the effects of prior social experience (with either a male or a female) on male investment in courtship and aggression under physiologically benign and challenging conditions (i.e., low versus high salinity). RESULTS: We found that males maintained a higher level of aggression towards a rival after a recent encounter with a female, compared to an encounter with a male, under low (but not high) salinity. In contrast, male investment in courtship behaviour was unaffected by either salinity or social experience. CONCLUSION: Together, our results suggest that male investment in aggression and courtship displays can differ in their sensitivity to environmental conditions and that not all reproductive behaviours are similarly influenced by the same environmental context.

A high aggression strategy for smaller males.

Male-male conflict is common among animals, but questions remain as to when, how and by whom aggression should be initiated. Factors that affect agonistic strategies include residency, the value of the contested resource and the fighting ability of the two contestants. We quantified initiation of aggression in a fish, the desert goby, Chlamydogobius eremius, by exposing nest-holding males to a male intruder. The perceived value of the resource (the nest) was manipulated by exposing half of the residents to sexually receptive females for two days before the trial. Resident male aggression, however, was unaffected by perceived mating opportunities. It was also unaffected by the absolute and relative size of the intruder. Instead resident aggression was negatively related to resident male size. In particular, smaller residents attacked sooner and with greater intensity compared to larger residents. These results suggest that resident desert goby males used set, rather than conditional, strategies for initiating aggression. If intruders are more likely to flee than retaliate, small males may benefit from attacking intruders before these have had an opportunity to assess the resident and/or the resource.

Both male and female identity influence variation in male signalling effort.

BACKGROUND: Male sexual displays play an important role in sexual selection by affecting reproductive success. However, for such displays to be useful for female mate choice, courtship should vary more among than within individual males. In this regard, a potentially important source of within male variation is adjustment of male courtship effort in response to female traits. Accordingly, we set out to dissect sources of variation in male courtship effort in a fish, the desert goby (Chlamydogobius eremius). We did so by designing an experiment that allowed simultaneous estimation of within and between male variation in courtship, while also assessing the importance of the males and females as sources of courtship variation. RESULTS: Although males adjusted their courtship depending on the identity of the female (a potentially important source of within-male variation), among-male differences were considerably greater. In addition, male courtship effort towards a pair of females was highly repeatable over a short time frame. CONCLUSION: Despite the plasticity in male courtship effort, courtship displays had the potential to reliably convey information about the male to mate-searching females. Our experiment therefore underscores the importance of addressing the different sources contributing to variation in the expression of sexually-selected traits.

Do male desert gobies compromise offspring care to attract additional mating opportunities?

Males often play a critical role in offspring care but the time and energy invested in looking after young can potentially limit their ability to seek out additional mating opportunities. Recent studies, however, suggest that a conflict between male parental effort and mating effort may not always be inevitable, especially if breeding occurs near the nest, or if parental behaviours are under sexual selection. Accordingly, we set out to experimentally investigate male care and courtship in the desert goby Chlamydogobius eremius, a nest-guarding fish with exclusive paternal care. Despite courtship occurring near the nest, we found that when egg-tending males were given the opportunity to attract additional females, they fanned their eggs less often, engaged in shorter fanning bouts, and spent more of their time outside their nests courting. Our findings highlight the importance of understanding the circumstances under which reproductive tradeoffs are expected to occur and how these, in turn, operate to influence male reproductive decisions.