Sirpa Kaunisto

New bedding site examination-based method to analyse deer ked (Lipoptena cervi) infection in cervids

Invasion of the deer ked (Lipoptena cervi), an ectoparasitic fly commonly found in cervids, has been rapid in Finland during the last four decades. As the distribution area of this species has expanded from the south towards the northern latitudes, the associated problems have become more evident. Various animals such as horses, cattle and especially reindeer have been reported to host this parasite. Moreover, in certain areas, the deer ked causes major inconveniences for humans potentially limiting recreational activities in forests. We wanted to study if deer ked parasitism and intensity of the infection in winter time could be detected by using visual examination of the snow on cervid bedding sites and by analysing biotic samples found from the bedding sites. Our results demonstrate that chronic deer ked infection causes reddish-brown snow discolouration (host tissue fluid and deer ked faeces) on the bedding sites to the extent that parasitism can be diagnosed. Hence, we suggest that deer ked infection prevalence and range expansion could be rapidly monitored using our new practical and reliable method. In the future, bedding site analyses will likely be useful in predicting and potentially preventing the negative effects of this ecologically and socio-economically important parasite.

Experiments on the ectoparasitic deer ked that often attacks humans; preferences for body parts, colour and temperature.

The deer ked (Lipoptena cervi) can fail in its host search. Host search fails when an individual deer ked irreversibly accepts a host unsuitable for its reproduction (e.g. a human) and drops its wings. In northern Europe, the main host of the deer ked is the moose (Alces alces). The deer ked is increasingly causing serious problems for humans (for example, causing deer ked dermatitis) and is considered a threat for the recreational use of forests. The adult deer ked flies in early and mid-autumn to search for a host. Our aims were: (i) to study whether there are ways to avoid deer ked attacks by wearing particular clothing, and (ii) to evaluate deer ked host choice. Using human targets, we explored the cues the deer ked uses for host selection. We studied which part of the host body deer keds target and if body colour and temperature affect their choice. In our experiments, deer keds landed more on dark and red clothing than on white clothing. Moreover, deer keds mostly attacked the upper body parts and preferred the back side of the body over the front side. Finally, deer keds preferred the warmest areas of the host.

High cold tolerance through four seasons and all free-living stages in an ectoparasite

Off-host stages of temperate parasites must cope with low temperatures. Cold tolerance is often highest in winter, as a result of diapause and cold acclimation, and low during the active summer stages. In some blood-feeding ectoparasites, offspring provisioning determines cold tolerance through all the non-feeding, off-host stages. Large size increases survival in the cold, but so far seasonal variation in within-female offspring size has not been associated with offspring cold tolerance. The deer ked (Lipoptena cervi) reproduces on cervids from autumn to spring. Newborn pupae drop off the host, facing frosts without any acclimation. We examined cold tolerance through 4 seasons and from birth to adulthood by means of short- and long-term frost exposure. We expected females to produce more tolerant offspring in winter than in spring. Large spring pupae survived prolonged frosts better than did small winter pupae. Thus more tolerant offspring were not produced when the temperature outside the host is at its lowest. Unexpectedly, the freezing points were -20 °C or below all year round. We showed that high cold tolerance is possible without acclimation regardless of life stage, which presumably correlates with other survival characteristics, such as the starvation resistance of free-living ectoparasites.

Northward invasion of the parasitic deer ked (Lipoptena cervi), is there geographical variation in pupal size and development duration?

The deer ked (Lipoptena cervi) is a common ectoparasite of cervids. During the last decades the species has rapidly invaded in northern Europe, especially in Finland, towards the north and increased its prevalence on the moose population. Consequently, during this rapid invasion the deer ked has faced more severe climatic conditions. We studied whether pupal size (measured as pupal weight) and pupal development duration of the deer ked varies along historical invasion zones and temperature zones towards north in Finland. Moreover, we explored possible size- and gender-dependent variation in pupal development duration. We divided wild-collected pupae in respect to their origin in two ways: (1) temperature zones (from south-west to colder north-east) and (2) invasion history (from early to late establishment). We reared pupae in the controlled laboratory conditions in identical temperature and light conditions. Pupal size decreased towards north and the smaller pupae developed faster. However, the results do not show differences in pupal size or developmental characteristics between the invasion zones. This supports the idea of rapid developmental plasticity of the deer ked and that not the invasion history but the current temperature regime determines the life history of the deer ked when invading towards a colder environment.

Early-life temperature modifies adult encapsulation response in an invasive ectoparasite.

Immunity of parasites has been studied amazingly little, in spite of the fact that parasitic organisms, especially the arthropod parasites, need immunity to survive their own infections to successfully complete life cycles. Long-term effects of challenging environmental temperatures on immunity have remained unstudied in insects and parasites. Our study species, the deer ked (Lipoptena cervi; Linnaeus 1758), is an invasive, blood-feeding parasitic fly of cervids. Here, it was studied whether thermal stress during the pupal diapause stage could modify adult immunity (encapsulation capacity) in L. cervi. The effect of either a low temperature or high temperature peak, experienced during winter dormancy, on encapsulation response of active adult was tested. It was found that low temperature exposure during diapause, as long as the temperature is not too harsh, had a favourable effect on adult immunity. An abnormal, high temperature peak during pupal winter diapause significantly deteriorated the encapsulation capacity of emerged adults. The frequency and intensity of extreme weather events such as high temperature fluctuations are likely to increase with climate change. Thus, the climate change might have previously unknown influence on host-ectoparasite interactions, by affecting ectoparasites immune defence and survival.

Summer time predation on the obligatory off-host stage of an invasive ectoparasite.

Predation can regulate populations and strongly affect invasion success of novel prey. The deer ked (Lipoptena cervi; Linnaeus 1758) is an invasive ectoparasite of cervids that spends a long period of its life cycle outside the host. Prior to this study, virtually nothing was known about natural summer time predation on the deer ked. We aimed to evaluate the magnitude of summer time predation on L. cervi pupae in different habitats and to identify potential predators. We conducted a set of field experiments, where we exposed L. cervi pupae to various ground-dwelling vertebrate and invertebrate predators. The loss of pupae was monitored for different predator guilds. Three habitats of the moose, the main host species, were studied: (1) moist heath forest; (2) dry, logged heath forest; and (3) moist meadow. The results indicate notable summer time predation on L. cervi pupae, and the pupal predation varied within and between habitats, being lowest in the meadow habitat. We found a positive correlation between pupal loss and abundance of the common lizard (Zootoca vivipara), harvestmen (Opiliones), ground spiders (Gnaphosidae) and Formicinae-ants. We conclude that summer time predation during the pupal phase can have a notable local importance for the L. cervi abundance.

Passive sinking into the snow as possible survival strategy during the off-host stage in an insect ectoparasite.

Abiotic and biotic factors determine success or failure of individual organisms, populations and species. The early life stages are often the most vulnerable to heavy mortality due to environmental conditions. The deer ked (Lipoptena cervi Linnaeus, 1758) is an invasive insect ectoparasite of cervids that spends an important period of the life cycle outside host as immobile pupa. During winter, dark-coloured pupae drop off the host onto the snow, where they are exposed to environmental temperature variation and predation as long as the new snowfall provides shelter against these mortality factors. The other possible option is to passively sink into the snow, which is aided by morphology of pupae. Here, we experimentally studied passive snow sinking capacity of pupae of L. cervi. We show that pupae have a notable passive snow sinking capacity, which is the most likely explained by pupal morphology enabling solar energy absorption and pupal weight. The present results can be used when planning future studies and when evaluating possible predation risk and overall survival of this invasive ectoparasite species in changing environmental conditions.