Many factors are involved in determining the latitudinal and altitudinal spread of the important tick vector Ixodes ricinus (Acari: Ixodidae) in Europe, as well as in changes in the distribution within its prior endemic zones. This paper builds on published literature and unpublished expert opinion from the VBORNET network with the aim of reviewing the evidence for these changes in Europe and discusses the many climatic, ecological, landscape and anthropogenic drivers. These can be divided into those directly related to climatic change, contributing to an expansion in the tick’s geographic range at extremes of altitude in central Europe, and at extremes of latitude in Scandinavia; those related to changes in the distribution of tick hosts, particularly roe deer and other cervids; other ecological changes such as habitat connectivity and changes in land management; and finally, anthropogenically induced changes. These factors are strongly interlinked and often not well quantified. Although a change in climate plays an important role in certain geographic regions, for much of Europe it is non-climatic factors that are becoming increasingly important. How we manage habitats on a landscape scale, and the changes in the distribution and abundance of tick hosts are important considerations during our assessment and management of the public health risks associated with ticks and tick-borne disease issues in 21st century Europe. Better understanding and mapping of the spread of I. ricinus (and changes in its abundance) is, however, essential to assess the risk of the spread of infections transmitted by this vector species. Enhanced tick surveillance with harmonized approaches for comparison of data enabling the follow-up of trends at EU level will improve the messages on risk related to tick-borne diseases to policy makers, other stake holders and to the general public.

/pdf/driving-forces-for-changes-in-geographical-distribution-of-1aaezoaq4a.pdf

Driving forces for changes in geographical distribution of Ixodes ricinus ticks in Europe

Allee effects: population growth, critical density, and the chance of extinction

The population ecology of ticks is fundamental to the spatial and temporal variation in the risk of infection by tick-borne pathogens. Tick population dynamics can only be fully understood by quantifying the rates of the demographic processes, which are influenced by both abiotic (climatic) factors acting on the free-living tick stages and biotic (host) responses to the tick as a parasite. Within the framework of a population model, I review methods and results of attempts to quantify (1) rates of tick development and the probability of diapause, (2) the probability of questing for hosts by unfed ticks, (3) the probability of ticks attaching to a host, and (4) tick mortality rates. Biologically, these processes involve the physiological and behavioural response of ticks to temperature, moisture stress and day length that result in specific patterns of seasonal population dynamics and host relationships. Temperate and tropical patterns will be illustrated with reference mostly to Ixodes ricinus and Rhipicephalus appendiculatus, respectively.

https://www.cambridge.org/core/services/aop-cambridge-core/content/view/S0031182004004925

Tick ecology: processes and patterns behind the epidemiological risk posed by ixodid ticks as vectors

During the last 30 years, there has been a continued increase in human cases of tick-borne encephalitis (TBE) in Europe, a disease caused by tick-borne encephalitis virus (TBEV). TBEV is endemic in an area ranging from northern China and Japan, through far-eastern Russia to Europe, and is maintained in cycles involving Ixodid ticks (Ixodes ricinus and Ixodes persulcatus) and wild vertebrate hosts. The virus causes a potentially fatal neurological infection, with thousands of cases reported annually throughout Europe. TBE has a significant mortality rate depending upon the strain of virus or may cause long-term neurological/neuropsychiatric sequelae in people affected. In this review, we comprehensively reviewed TBEV, its epidemiology and pathogenesis, the clinical manifestations of TBE, along with vaccination and prevention. We also discuss the factors which may have influenced an apparent increase in the number of reported human cases each year, despite the availability of effective vaccines.

Tick-borne Encephalitis Virus - A Review of an Emerging Zoonosis

The statistical aggregation of parasites among hosts is often described empirically by the negative binomial (Poisson-gamma) distribution. Alternatively, the Poisson-lognormal model can be used. This has the advantage that it can be fitted as a generalized linear mixed model, thereby quantifying the sources of aggregation in terms of both fixed and random effects. We give a worked example, assigning aggregation in the distribution of sheep ticks Ixodes ricinus on red grouse Lagopus lagopus scoticus chicks to temporal (year), spatial (altitude and location), brood and individual effects. Apparent aggregation among random individuals in random broods fell 8-fold when spatial and temporal effects had been accounted for.

Analysis of aggregation, a worked example: numbers of ticks on red grouse chicks

1. Activity in Ixodes ricinus was studied by laying down newly emerged ticks in natural clumps of vegetation from which all hosts (with the possible exception of ‘mice’) were excluded. After a period of quiescence the ticks climbed to the ‘active’ position at the vegetation tips. Activity was assessed either by simple observation (in adults which were marked individually) or by ‘brushing’ the vegetation with the hands (nymphs and larvae). 52 % of the adults and 44 % of the nymphs were recovered.2. The seasonal activity behaviour was followed in three series of ticks set out during May, July and October 1945. The results were used in interpreting the time relations of seasonal activity in ‘wild’ populations. The timing of the activity cycle appears to be largely determined by the availability of unfed ticks rather than by the prevailing meteorological conditions.3. The diurnal pattern of behaviour in active ticks was closely examined during a 24 hr. period. Many remained at the tips without moving. The others began or ended a phase of activity by night or by day; but among these, the proportion ending a phase at night was significantly greater. This is the main reason for the previously reported diurnal fluctuation of activity in a tick population.4. Two hundred and seven adult ticks spent, on an average, a total of 9 days at the vegetation tips (individual limits 1 and 54 days). This activity was spread over an ‘active period’ (interval between first and last appearances) averaging 30 days. Very active ticks rarely remained continuously at the tips. The mean number of visits to the tips was 4 (limits 1 and 19), each lasting for an average of 2·5 days. Where long spells of activity alternate with short periods of quiescence, the behaviour is probably regulated by the humidity reaction, the quiescent phase providing an opportunity for restoring the depleted water balance.5. Newly moulted adult ticks possess reserves of fat adequate for many months of quiescence. During activity these reserves are exhausted in a few weeks or even days. Duration of survival mainly depended on whether the onset of activity was immediate or delayed. Some adults remained quiescent for 1 year before becoming active for the first time.6. Several types of orientations were observed in the field. In attaining the position at the tips favourable for encountering a host, the gravity response (upward-turning near the tip) is of major importance. The tick avoids wind and direct sunlight by sheltering behind its supporting stem. In sensing the approach of a ‘host’ (the observer's finger) the perception of eddies of warm air is particularly significant. The response (questing, then orientation) is elicited much less readily if the stimulus is applied from the leeward.

https://www.cambridge.org/core/services/aop-cambridge-core/content/view/S0031182000084031

The seasonal and diurnal activities of individual sheep ticks (Ixodes ricinus L).

Since it is concerned with filling in certain details of a picture, long since sketched in outline, the new work in this paper permits only a somewhat disjointed summary. From a study of Ixodes ricinus L. in nature, the main new facts emerging are: (I) Unfed adults in a deep vegetation layer such as presented by rough hill or moorland pasture dominated by bents, heather or bracken. (i) Practically all inactive unfed ticks are in the underlying mat, and nearly all of these in the upper portion of the mat. This distribution does not change, summer or winter. (ii) When the tick is active its progression is practically confined to the vertical, i.e. between its niche in the mat and the vegetation tips immediately above. Random undirected movements due to certain circumstances may achieve a 0–8 in. (average 2 in.) change of position in the horizontal plane; but there is apparently no ruling urge to move in the horizontal plane. (iii) The unfed tick comes to the vegetation tips to await a host for only a limited time during the 3-month activity season. Commonly this amounts to about five periods of 4–5 days each if no host is forthcoming. Between the periods of activity it returns to the upper mat. If it stayed constantly at the tips its survival time would be much shorter than normal. (iv) From nymphs engorged in spring, adults emerge in autumn but remain inactive for a considerable period. About half of them may perish within the first 100 days after emergence. Survivors become active during the following March–June. Effective life ends when the tick, if still unfed, finally stops coming to the vegetation tips, the only position from which a host can normally be achieved. After emergence, the effective life of the unfed adult deprived of a host is less than one year; and the actual life is not much more than one year at best. Thus adults not finding a host in spring will be ineffective, i.e. die without reproducing before the next activity season.

The ecology of the sheep tick, Ixodes ricinus L.; microhabitat economy of the adult tick.

Although three other Ixodes species were found, Ixodes ricinus L., when it occurs, is by far the most important tick parasite of wild and domestic animals on hill and moorland grazings in northern England. In the following, ‘tick’ or ‘ticks’ refer to I. ricinus.The previous list of British mammal hosts, wild and domestic, to the tick (any stage) is largely confirmed, and increased approximately from twenty-three to twenty-nine species; and birds, similarly, from twenty-one to thirty-nine species. The preponderance of small mammals and birds among the twenty-four new-recorded host species confirms the impression that previous workers tended to confine their attentions to the larger animals. The host list is still incomplete, and the suggestion that the tick will parasitize any bird or mammal it may meet is strongly reinforced.No host occurs below the reptile level among vertebrates; and among British reptiles, only the common lizard is a host—rarely. There appear to be no hosts among invertebrates.All the farm and domestic mammals from horse down to cat are hosts to female ticks, but poultry have never been recorded—probably because they do not much frequent the tick-infested pastures.Only the larger wild mammals and birds are hosts to female ticks. In northern England these include eight mammals and three birds. For Britain as a whole two other mammals are known, while a further four mammals and one bird are likely hosts.With or without the inclusion of domestic animals, proportionately more mammal than bird species are hosts to female ticks. This is because there are more mammal than bird species large enough to be hosts.All the recorded British hosts, except shrew and possibly mole, are hosts in some degree to nymphs, and all, without exception, are hosts to larvae.A certain amount of self-deticking is practised by birds, and very probably by some of the smaller mammals. Birds never have ticks attached within beak-reach.

https://www.cambridge.org/core/services/aop-cambridge-core/content/view/S0031182000083736

The ecology of the sheep tick, Ixodes ricinus L. Host relationships of the tick: Part 2. Observations on hill and moorland grazings in northern England

Spatial distribution is understood to connote distribution of density (including zero density) over the land surface at a given time. Combining fact and conjecture, the following is a theory of spatial distribution of Ixodes ricinus in Britain. The tick passes practically the whole of its 3-year life span on the ground, spending a total of only 3 weeks on hosts. A considerable amount of vegetational cover is necessary for efficient survival on the ground; the more important limiting circumstances in lesser cover seem to be humidity (conditioned by temperature) in summer, perhaps temperature (alone) in winter, and the activity of predators. A host is necessary for mating as well as food purposes. Though small at best, the tick's chances of achieving a host are greater the more hosts on the ground. In practice this means the more sheep, since the latter are by far the main hosts. For the most part, spatial distribution to-day can be explained adequately in terms of interplay between amount of ground-cover and of host-potential, with cover the master factor. Thus poor cover keeps the heavily stocked lowland pastures free of permanent tick population even when opportunity of colonization is offered. The tick is confined to the hills and moorlands with their deep vegetation layers. There, within limits, it is more numerous on a pasture the more universal the distribution of adequate cover (because it drops at random from the host) and the higher the host-potential. Among hill and moorland pastures, however, there are some cases where, despite ample cover and hostpotential, either (1) no ticks are present, or (2) the tick population is lower than it ought to be. These cases can be explained on grounds of lack of opportunity for colonization (1); or colonization so recent that population has not attained the level of ‘dynamic equilibrium’ with the environment (2). The question of ‘spread’ (colonization) is discussed fully in the text. The sheep flock is the main agent in spreading ticks. Spread is comparatively slow and relatively small in extent nowadays. There has very probably been considerable increase both in tick population densities and in infested areas since the advent of intensive hill and moorland grazing with farm stock some centuries ago.

The ecology of the sheep tick, Ixodes ricinus L.; spatial distribution.

In three experiments, the chief hosts, sheep, were withheld from a grazing until 0–3 weeks after the normal spring peak of female tick activity. This did not give rise to any appreciable prolongation of female tick activity into the normal summer offseason. More than half the female ticks actually available during the entire spring season were unfed when activity came to an end. This was due to the withholding of hosts. Hence that aspect of the ‘two-brood’ theory which claims that activity ends in spring, because all the available ticks are fed, is untenable as the complete explanation of bimodal annual activity. Increase in sheep-stocking density, over the normal for a grazing, lowers the average infestation per sheep for that season at least. The experimental data suggests the following: (i) Delay in stocking a grazing with sheep results in a slightly higher-than-normal population of active female ticks which is fairly quickly absorbed after reintroduction of sheep. (ii) A varying proportion of the tick population becomes active each week throughout the tick season, increasing up to the peak and decreasing thereafter. (iii) The average duration of individual activity in a natural population of female ticks is probably not more than 1 month, perhaps less. Practical applications of the findings to hill sheep farming are indicated.

https://www.cambridge.org/core/services/aop-cambridge-core/content/view/S0031182000012117

A. Milne

Papers

The seasonal and diurnal activities of individual sheep ticks (Ixodes ricinus L).

The ecology of the sheep tick, Ixodes ricinus L.; microhabitat economy of the adult tick.

The ecology of the sheep tick, Ixodes ricinus L. Host relationships of the tick: Part 2. Observations on hill and moorland grazings in northern England

The ecology of the sheep tick, Ixodes ricinus L.; spatial distribution.

The ecology of the sheep tick, Ixodes ricinus L. Host availability and seasonal activity.