Changes in Habitat Suitability for the Tick Ixodes ricinus (Acari: Ixodidae) in Europe (1900–1999)

Climatic changes may lead to drastic changes in the distribution of arthropods important in human health. We tracked changes in habitat suitability for the tick Ixodes ricinus in Europe from 1900 to 1999, using a geographically extensive gridded climate data set. For the whole period, 52% of the territory was always unsuitable for the tick. In the grid, 6.11% of the cells were classified as having a deterministic drift with positive trend and 7.4% as deterministic drift with a negative trend. A total of 17.25% of cells were classified as exhibiting a random walk behavior, with a trend to increase of habitat suitability (9.57%) or decrease (7.68%). Zones of deterministic trend extend into most of Ireland and parts of the United Kingdom and France. Total and summer rainfall primarily drive changes in habitat suitability in these sites. Areas of random walk are common in Scandinavia, central Europe, and the Balkans, with summer rainfall and temperature largely directing the changes. Sites of reported increased abundance of I. ricinus coincided with areas of increased habitat suitability over the last 20–30 years, but this feature showed a long-term random walk negative trend. Habitat suitability for I. ricinus remains relatively stable in Europe, with no sites showing permanent changes in habitat suitability (negative to sustained positive or vice versa). However, some zones in the continent showed a clear trend to increase or decrease.     

An experimental test to compare potential and realised specificity in ticks with different ecologies

The majority of studies on ecological specialisation rely on data reflecting realised specificity, without considering species’ potential specificity. Most species of ticks, a large family of hematophagous ectoparasites, have a narrow host range in nature, but it is unclear whether this is due to host-driven adaptations or other processes (such as off-host abiotic environment). We investigated the potential specificity of two tick species with contrasting ecology by infesting three avian host species that occur in the same off-host macrohabitat but are unequally infested by the ticks in nature (i.e. have contrasting realised specificity). The endophilic specialist tick Ixodes arboricola resides inside the hosts’ nest and has high realised host specificity, whereas the exophilic generalist tick I. ricinus encounters hosts in the field and has very low realised specificity. As hosts, we used great tits (frequently infested by both tick species), blackbirds (frequently infested by I. ricinus but never by I. arboricola) and great spotted woodpeckers (no ticks of either species have been reported). If realised specificity is constrained by host-driven adaptations there should be no differences between potential and realised specificity, whereas if realised specificity is constrained by other processes potential specificity and realised specificity should be different. We found that attachment rates and weight during feeding of I. arboricola were lower on blackbirds than on great tits, whereas there were no such differences for I. ricinus. No ticks of either species attached to woodpeckers. These results indicate that realised host specificity of ticks is, at least partially, constrained by host-driven adaptations. This specificity therefore strongly depends on the ticks’ encounter rates with particular host types, which are affected by the ticks’ off-host ecological requirements, behaviour and life-history characteristics.     

Forecasting next season’s <Emphasis Type="Italic">Ixodes ricinus</Emphasis> nymphal density: the example of southern Germany 2018

The castor bean tick, Ixodes ricinus (L.) (Ixodida: Ixodidae), is the principal vector of pathogens causing tick-borne encephalitis or Lyme borreliosis in Europe. It is therefore of general interest to make an estimate of the density of I. ricinus for the whole year at the beginning of the tick season. There are two necessary conditions for making a successful prediction: a long homogeneous time series of observed tick density and a clear biological relationship between environmental predictors and tick density. A 9-year time series covering the period 2009–2017 of nymphal I. ricinus flagged at monthly intervals in southern Germany has been used. With the hypothesis that I. ricinus density is triggered by the fructification of the European beech 2 years before, the mean annual temperature of the previous year, and the current mean winter temperature (December–February), a forecast of the annual nymphal tick density has been made. Therefore, a Poisson regression model was generated resulting in an explained variance of 93.4% and an error of \(\hbox {RMSE} = 21\) ticks per \(100\,\hbox {m}^2\) (annual \(\hbox {MEAN} = 260\) collected ticks/\(100\,\hbox {m}^2\)). An independent verification of the forecast for the year 2017 resulted in 187 predicted versus 180 observed nymphs per \(100\,\hbox {m}^2\). For the year 2018 a relatively high number of 443 questing I. ricinus nymphs per \(100\,\hbox {m}^2\) is forecasted, i.e., a “good” tick year.     

Predicting the distribution of <Emphasis Type="Italic">Encephalartos latifrons</Emphasis>, a critically endangered cycad in South Africa

This study evaluates how a modelling approach to determine areas of suitable habitat for the Critically Endangered Albany cycad Encephalartos latifrons can assist in systematic conservation planning for this and other rare and threatened cycads. A map distinguishing suitable from unsuitable habitat for E. latifrons was produced and important environmental predictors (climate, geology, topography and vegetation) influencing the suitable habitat were estimated. The maximum entropy (MaxEnt) modelling technique was chosen for this study as it has consistently performed well compared with alternative modelling methods and is also an appropriate model choice when the sample size is small and locality records are relatively few. Predicted habitat suitability showed that some locations chosen for translocation and restoration of E. latifrons specimens are not suitable. This revealed that modelling suitable habitat can guide relocation and regeneration of E. latifrons and perhaps other threatened cycads with restricted distributions and few locality records. The species distribution model constructed for E. latifrons is the first reported habitat model for a Critically Endangered cycad in South Africa. The results may be incorporated into conservation planning and structured decision-making about translocations and restoration programmes involving vulnerable cycads, which are among the most threatened organisms globally.     

A species distribution model for pine marten (<Emphasis Type="Italic">Martes martes</Emphasis>) in the least forested region of Europe

Species distribution modelling is a useful technique that provides data on factors that can influence a species range, identify high suitability areas and model future scenarios. The pine marten (Martes martes) has undergone major historical declines in abundance and distribution in Northern Ireland, similar to that which has occurred throughout its range. Currently, the species is in a phase of range expansion in Northern Ireland, in what is the least forested landscape in Europe. To assess the suitability of this environment for pine marten re-establishment, presence only distribution data combined with landcover data at a 10-km scale were used in a species distribution modelling study using Maxent. The results indicated that approximately 32% (4500 km²) of the land area of Northern Ireland had a high probability of pine marten occurrence. Pine marten distribution was positively associated with the extent of conifer forest landcover types, which also had the highest single attribute contribution to the model. Landcover types that were negatively associated with pine marten distribution included the extent of open, dwarf and urban areas.     

Ticks in the ecotone: the impact of agri-environment field margins on the presence and intensity of Ixodes ricinus ticks (Acari: Ixodidae) in farmland in southern England

The present study aimed to assess whether agri-environment field margins provide a habitat for the sheep/deer tick Ixodes ricinus. Field studies were conducted in arable farmland in southern England in both extant and newly constructed field margins. The presence and intensity (i.e. the mean number of nymphs per transect, excluding zeros) of questing nymphs and adult I. ricinus were compared between field margins with three adjacent habitats: woodland, hedgerow and arable land. The presence and intensity of ticks within a field margin was also compared between three ecozones: the ecotone, the margin and the crop. It was found that field margins do support I. ricinus, although the intensity of ticks was associated with field margins with adjacent woodland, with a higher tick intensity along the ecotonal ecozone, compared with the rest of the margin or the crop edge. The presence of a hedge also increased the likelihood of finding questing nymphs in a field margin compared with a margin adjacent to arable land. This effect, however, was less pronounced than in field margins with adjacent woodland. The provision of footpaths within the margin (at least 1–2 m from the ecotone), or on the edge of the crop where paths run next to woodland known to be an important tick habitat, could be promoted to minimize tick exposure. In addition, based on the results of the present study, raising awareness that walking alongside woodlands also constitutes a tick risk could be promoted.     

Assessment of Habitat Fragmentation and Corridors for an Isolated Subspecies of the Sichuan Golden Snub-Nosed Monkey, <Emphasis Type="Italic">Rhinopithecus roxellana hubeiensis</Emphasis>

Understanding habitat quality and landscape connectivity and exploring corridors connecting habitat patches are crucial for conservation, particularly for species distributed among isolated populations. The Sichuan golden snub-nosed monkey, Rhinopithecus roxellana, is an Endangered primate species endemic to mountainous forests in China. Its easternmost distribution lies in the Shennongjia area, which harbors an isolated subspecies, R. roxellana hubeiensis. Unfortunately, it has experienced significant habitat loss, fragmentation, and dramatic population decline in recent decades, primarily due to increased human disturbance. To quantify habitat quality, identify suitable habitat patches, and detect possible linkages among these patches for R. roxellana hubeiensis, we conducted habitat suitability assessments and landscape connectivity analyses in the Shennongjia area based on a set of environmental factors. We created a habitat quality model and a movement cost surface for the Shennongjia area based on a habitat suitability index, graph theory, expert knowledge, field experience, and information from the literature. Our results show that suitable habitat for R. roxellana hubeiensis in Shennongjia is fragmented and limited, and that this is particularly true for highly suitable habitats. We detected six core habitat patches and six least-cost paths and corridors. Our study does not provide accurate distributions of the monkeys and their habitat use. However, it identifies the most feasible and traversable habitats and corridors, which should be conservation priorities for this subspecies, and provides valuable guidance for reevaluating habitat conservation plans.     

Possible expansion of Ixodes ricinus in the United Kingdom identified through the Tick Surveillance Scheme between 2013 and 2020

The tick Ixodes ricinus (Ixodida: Ixodidae, Linnaeus) is the main vector of several pathogens including Borrelia burgdorferi s.l. (agent of Lyme borreliosis) and tick-borne encephalitis virus. Its distribution depends on many factors including suitable habitat, climate and presence of hosts. In this study, we present records of I. ricinus bites on humans, dogs (Canis lupus familiaris; Carnivora: Canidae, L.) and cats (Felis catus; Carnivora: Felidiae, L.) in the United Kingdom (UK) obtained through the Tick Surveillance Scheme between 2013 and 2020. We divided the UK into 20 km x 20 km grids and 9.2% (range 1.2%–30%) of grids had at least one record every year since 2013. Most regions reported a yearly increase in the percentage of grids reporting I. ricinus since 2013 and the highest changes occurred in the South and East England with 5%–6.7% of new grids reporting I. ricinus bites each year in areas that never reported ticks before. Spatiotemporal analyses suggested that, while all regions recorded I. ricinus in new areas every year, there was a yearly decline in the percentage of new areas covered, except for Scotland. We discuss potential drivers of tick expansion, including reforestation and increase in deer populations.     

Occupancy modeling of <Emphasis Type="Italic">Parnassius clodius</Emphasis> butterfly populations in Grand Teton National Park,Wyoming

Estimating occupancy patterns and identifying vegetation characteristics that influence the presence of butterfly species are essential approaches needed for determining how habitat changes may affect butterfly populations in the future. The montane butterfly species, Parnassius clodius, was investigated to identify patterns of occupancy relating to habitat variables in Grand Teton National Park and Bridger-Teton National Forest, Wyoming, United States. A series of presence–absence surveys were conducted in 2013 in 41 mesic to xeric montane meadows that were considered suitable habitat for P. clodius during their flight season (June–July) to estimate occupancy (ψ) and detection probability (p). According to the null constant parameter model, P. clodius had high occupancy of ψ?=?0.78?±?0.07 SE and detection probability of p?=?0.75?±?0.04 SE. In models testing covariates, the most important habitat indicator for the occupancy of P. clodius was a strong negative association with big sagebrush (Artemisia tridentata; β = ??21.39?±?21.10 SE) and lupine (Lupinus spp.; β?=???20.03?±?21.24 SE). While P. clodius was found at a high proportion of meadows surveyed, the presence of A. tridentata may limit their distribution within montane meadows at a landscape scale because A. tridentata dominates a large percentage of the montane meadows in our study area. Future climate scenarios predicted for high elevations globally could cause habitat shifts and put populations of P. clodius and similar non-migratory butterfly populations at risk.     

The geographic distribution and ecological preferences of the American dog tick,Dermacentor variabilis (Say), in the U.S.A.

Equine piroplasmosis (EP), caused by two parasitic organisms, Theileria equi and Babesia caballi, is a tick‐borne disease of recent concern in horses in the U.S.A. Outbreaks of EP have been detected in Florida, Missouri, Kansas and Texas. In 2009, EP transmission in Texas occurred through the adults of two tick species, Amblyomma mixtum [formerly known as Amblyomma cajennense (Fabricius, 1787)] Koch (Ixodida: Ixodidae) and Dermacentor variabilis (Say) (Ixodida: Ixodidae), the American dog tick (ADT). In this study, we developed a continent‐scale map for the distribution of the EP vector species D. variabilis, using a presence‐only modelling approach to assess the habitat preferences of this tick. We used identification records from our tick geodatabase of locations in which the presence of the ADT had been noted. The potential distribution of the ADT in the U.S.A. was estimated from environmental factors using the maximum entropy approach based on localities in which there is a high probability of occurrence according to habitat suitability. Elevation and temperature were found to be biologically significant environmental variables influencing the presence of this tick species. Properly designed and constructed probability surfaces using maximum entropy offer one useful approach to the mapping of distribution ranges of tick species based on suitable habitat in the U.S.A.     

Projecting present and future habitat suitability of ship-mediated aquatic invasive species in the Canadian Arctic

A rise in Arctic shipping activity resulting from global warming and resource exploitation is expected to increase the likelihood of aquatic invasive species (AIS) introductions in the region. In this context, the potential threat of future AIS incursions at a Canadian Arctic regional scale was examined. Habitat suitability under current environmental conditions and future climate change scenarios was projected for a subset of eight potential invaders ranked as having a high risk of establishment in the Canadian Arctic based on dispersal pathways/donor regions, biological attributes and invasion history: (1) Amphibalanus improvisus, (2) Botrylloides violaceus, (3) Caprella mutica, (4) Carcinus maenas, (5) Littorina littorea, (6) Membranipora membranacea, (7) Mya arenaria and (8) Paralithodes camtschaticus. Habitat modelling was performed using MaxEnt based on globally known native and non-native occurrence records and environmental ranges for these species. Results showed that under current environmental conditions the habitat is suitable in certain regions of the Canadian Arctic such as the Hudson Complex and Beaufort Sea for L. littorea, M. arenaria and P. camtschaticus. Under a future climate change scenario, all species showed poleward gains in habitat suitability with at least some regions of the Canadian Arctic projected to be suitable for the complete suite of species modelled. The use of these models is helpful in understanding potential future AIS incursions as a result of climate change and shipping at large spatial scales. These approaches can aid in the identification of high risk regions and species to allow for more focused AIS monitoring and research efforts in response to climate change.     

Woodland biodiversity management as a tool for reducing human exposure to Ixodes ricinus ticks: A preliminary study in an English woodland

This paper presents preliminary findings towards developing a UK‐specific approach to reducing public exposure to woodland questing Ixodes ricinus tick populations by harnessing existing biodiversity‐enhancing woodland ride (i.e., linear non‐wooded herbaceous habitat either side of track within woodland) management strategies. This preliminary study in an English woodland firstly assesses whether ecological and environmental factors determine presence and density of questing Ixodes ricinus along woodland rides. Secondly, it sets these findings in the context of woodland ride management guidelines in England in order to understand what impact ride management strategies might have on numbers of questing ticks and tick survival. Nymph and adult I. ricinus presence and abundance were modelled in relation to relevant microclimate and ecological parameter variables. Predictor variables for increased questing nymph abundance included ride orientation, mat depth, occurrence of bracken/bramble and animal tracks, ride/path width, and sward height. Ticks thrive in the ecotonal habitat of a woodland ride, therefore we urge woodland managers to consider the impact of their ride management on ticks and human exposure to ticks. Possible recommendations for mitigating questing I. ricinus in line with biodiversity management guidelines rides are discussed in this paper and include seasonal mowing regimes, management of mulch/mat, and bracken/bramble management through use of scalloped ride edges.     

Efficiency of flagging and dragging for tick collection

It is acknowledged that data from field studies on tick ecology might be biased by collection methods, but actually comparative studies are still limited. Herein we assessed whether the efficiency of flagging and dragging varies according to tick developmental stage, species, season and habitat. Ticks were collected in three sites bordered by an oak forest. The abundance of ticks collected by each collection method varied according to tick species, developmental stage, season, and habitat. Flagging was in general more efficient in collecting adult ticks, especially in spring and winter. Females were more frequently collected by flagging in the meadow and grassland habitats and males in the man-made trail. Flagging collected significantly more adults of Dermacentor marginatus, Hyalomma marginatum, Haemaphysalis inermis and Ixodes ricinus. Flagging was more efficient in collecting D. marginatus and I. ricinus in spring, and H. inermis and I. ricinus females in both spring and winter. In summer and autumn tick abundances were generally similar, with the exception of D. marginatus female in autumn. Flagging was more efficient in collecting D. marginatus adults in the meadow habitat and in the man-made trail, and I. ricinus adults in the meadow and grassland habitats. Dragging was more efficient in grassland for R. turanicus. Our results suggest that variations in terms of collection method performance are associated to factors linked to tick behaviour, habitat characteristics, and climate. Field studies employing these collection methods should take this into account to avoid misleading conclusions about tick population dynamics and tick-borne pathogen transmission risk.     

Will climate change affect ectoparasite species ranges?

Aim  Over the next 100 years, human-driven climate change and resulting changes in species occurrences will have global impacts on biodiversity, ecosystem function, and human health. Here we examine how climate change may affect the occurrences of tick species in Africa and alter the suitability of habitat outside Africa for African ticks.
Location  Africa and the world.
Methods  We predicted continental and global changes in habitat suitability for each of 73 African tick species, using multiple regression models in different climate change scenarios that cover a wide range of uncertainty.
Results  Global habitat suitability improves for nearly all tick species under each of a representative range of eight climate change scenarios. Depending on the scenario, African tick species experience an average increase in global habitat suitability of between 1 million and 9 million square kilometres between 1990 and 2100.
Main conclusions  The potential for successful translocations of ticks and their pathogens from Africa to the rest of the world is likely to increase over the next 100 years. Although the general trend is one of range expansion, there are winners and losers among tick species in each scenario, suggesting that tick community composition will be disrupted substantially by climate change. If this is also typical of other invertebrates, then climate change will disrupt not only the geographic location of communities but also their structure. Changes in tick communities are also likely to influence tick-borne pathogens.     

Blood-sucking ticks (Acarina,Ixodoidea) as an essential component of the urban environment

The problem of urban ticks has arisen from the increased rate of urbanization since WWII. Expansion of municipal boundaries encompasses adjacent territories, so that large areas of wilderness together with all their inhabitants get incorporated into city limits. Current strategies of biodiversity conservation include the creation of green corridors and other forms of connectivity between wilderness and urban areas as well as between green patches within cities. All this allows various mammals and birds to migrate from their native habitats into and between various parts of the cities and to establish permanent urban populations. Medium-sized and larger animals provide adult ticks with blood meal, thus creating suitable conditions for the establishment and persistence of tick populations. Independent tick populations can exist in urban forests, parks, private properties, old cemeteries, etc. Over the last decades, the tick populations that originated from those in natural habitats around the cities have become a permanent component of urban fauna. Among such ticks, the castor bean tick Ixodes ricinus is the most important species for European cities, while the deer tick I. scapularis is of great significance for the East Coast of the United States. The taiga tick I. persulcatus is the most important species for the Russian cities and towns within its range. All these and some other ticks aggressively attack humans and their pets inside cities. The tick species especially well adapted to urban life are those which can live and reproduce in buildings. The brown dog tick Rhipicephalus sanguineus (family Ixodidae), the pigeon ticks from the reflexus group of the genus Argas, and Ornithodoros ticks (family Argasidae), which form urban and semi-urban populations, are the main urban tick pests and vectors. House infestation by ticks can lead to human infection with tick-borne pathogens or severe allergic reactions. Some tick hosts, mammals as well as birds, maintain tick-transmitted pathogens and serve as competent reservoir hosts. Urban populations of these animals can participate in the circulation of some pathogens within cities. Thus, the enlargement of urban green areas followed by their population by wild mammals and birds create good opportunities for the establishment of urban populations of tick vectors with the resulting threat to the health of urban dwellers and their pets. At the same time, our understanding of the real scope and complexity of the problem of urban ticks is far from being sufficient. Finding the ways of protecting the environment without increasing the risk to human health in modern cities is a pressing and challenging problem of our time.     

Differential diagnosis of <Emphasis Type="Italic">Ixodes ricinus</Emphasis> and <Emphasis Type="Italic">Ixodes persulcatus</Emphasis>: nymphs and larvae

We developed a method for differential diagnosis of nymphs and larvae of sheep (Ixodes ricinus (L.)) and taiga (I. persulcatus Sch.) ticks (Parasitiformes: Ixodidae) which allows to identify live material in the field.     

Demographic effects of prolonged drought on a nascent introduction of a semi-aquatic snake

Faced with a nascent introduction of a non-native species, conservationists need to quickly determine how a population performs in its new environment. Although correlative models can predict environmental suitability at a coarse scale, they often neglect short-term climatic variability, instead relying on long-term averages. Accurately projecting the fate of any particular introduction requires demographic data on how a population responds to a novel environment. The recent introduction of watersnakes (genus Nerodia) poses a risk to California’s already imperiled aquatic vertebrate fauna. Despite inhabiting a seemingly suitable climate, a non-native Nerodia sipedon population in central California is likely to have been affected by a prolonged extreme drought from 2012 to 2015. We studied the only known population of N. sipedon in California for 3 years from 2013 to 2015, and estimated its abundance and annual survival. Its abundance declined from a peak of 218 (95% Credible Interval 149–313) in August 2013 to 97 (80–119) in July 2015. Annual survival of N. sipedon from 2013 to 2014 (0.23, 0.13–0.39) and from 2014 to 2015 (0.29, 0.18–0.41) was lower than survival estimates reported from native populations. Snake body condition, the abundance of large adult females, and prey availability all declined throughout the study. We conclude that the population of N. sipedon declined from 2013 to 2015, likely due to decreasing habitat and prey availability from the prolonged regional drought. This study highlights the importance of the effect of climatic extremes on the trajectory of introduced populations in a novel environment.     

New insights into the distribution and conservation status of the Golden-White Tassel-Ear Marmoset <Emphasis Type="Italic">Mico chrysoleucos</Emphasis> (Primates,Callitrichidae)

Among the 13 Mico species recognized by the IUCN Red List of Threatened Species, six are listed as “Data Deficient”. The geographic range of most of the Mico species has been estimated from only a few records. We report new localities and the geographic extension of Mico chrysoleucos. In addition, we confirmed the presence of the species in two distinct protected areas. We modeled the habitat suitability of M. chrysoleucos using the maximum entropy method and including new records obtained by the authors in the state of Amazonas, Brazil. From the total area of occurrence calculated for the species, 22.8% is covered by protected areas and indigenous lands. The annual mean deforestation rate estimated between 2000 and 2015 was 2.95%, and the total area deforested by 2015 was 3354 km² or 8.6% of the total distribution limits of the species. The habitat lost between 2000 and 2015 was 3.2% (1131 km²) of the total potential distribution, while the habitat loss area legally protected was 31 km², and the habitat loss in settlements was equal to 691 km². Our results extend the geographic distribution of the species about 100 km farther south, with the Maracanã River being a possible geographic barrier for the species. The significantly low rate of habitat loss inside protected areas and indigenous land, when compared to unprotected areas, points out the importance of these areas to M. chrysoleucos conservation. The species is relatively wide-ranging, legally protected, and resilient to regional anthropic threats. However, the hydroelectric schemes and the improvement of the road system in southern Amazonia pose an imminent threat to the species.     

Evaluation of the role of the national parks for Persian leopard (<Emphasis Type="Italic">Panther pardus saxicolor</Emphasis>, Pocock 1927) habitat conservation (case study: Tandooreh National Park,Iran)

Having knowledge of the habitat requirements of top predators is important for their conservation and also for the stability of wildlife communities since they are keystone species of ecosystems. This study aims to predict suitable habitats for the Persian leopard (Panther pardus saxicolor, Pocock 1927) in Tandooreh National Park, Iran. This species is the largest leopard subspecies in the Middle East and is at high risk of extinction. Presence points for the species have been obtained from field data, literature review, atlas data, and IUCN records. Variables related to human disturbance, terrain, land use/land cover, and the leopards’ prey were used to predict the habitat selection of the Persian leopard by employing binary logistic regression. Our model predicted that the presence of Persian leopards is restricted to the southern, north-western, and western areas of the park which are under severe fragmentation threat. Moreover, leopards avoided areas near human development such as roads. We have also found that the availability of prey such as Capra aegagrus, Ovis vignei, and Ovis orientalis is one of the important parameters to predict habitat suitability for leopards. Our model provides a suitable tool for national park managers to identify biodiversity hotspots since leopards are often associated with high biodiversity.