P. W. This pattern suggests that ticks acquire B. burgdorferi from reservoir hosts rather than transovarian transmission. Yang A. R. Talbert Finally, we identified counties where surveillance records indicate that Ixodes ticks are not established but at least two ensemble members predicted suitable habitat. Larvae hatch from fertilized eggs in late spring and feed once for 2 or more days in midsummer. D. For each variable, the value of deviance explained is parenthetically accompanied by the rank of this value in the list of values for all of the 68 originally considered predictor variables, arranged in descending order (i.e., a rank of 1 indicates the highest value). . W. S. b Candidate variables (predictor set 2): Bio15, Bio19, Bio1, Bio17, Bio8, Vp10, GDD2Cum, PercForest. J. It is the principal vector of Lyme disease in that region. 2003 ) using similar county-level data on the distribution of I. scapularis compiled nearly two decades ago ( Dennis et al. (Right column) Maps depicting the predicted distribution of I. pacificus by each of the individual optimized models: GLM, MARS, Maxent, and RF, respectively. 1998 ) similar to the data used in our models, Brownstein et al. The white-tailed deer is the preferred host of the adult I. scapularis. Khatchikian Across models, these variables combined explained between 41–86% of variation in suitability. Dippel There was a strong positive linear association between vapor pressure in February and habitat suitability, while the association between average days per month with snow cover and habitat suitability was weakly negative. Kunz P. M. Roe Hickling We used average climate, elevation, and land cover values for each county as predictors in our habitat models. Adult I. scapularis has a narrower host range, with a preference for deer. K. J. We also conducted a search to ensure that papers using the junior syno-nym “Ixodes dammini” rather than Ixodes scapularis … Lacombe T. J. The I. pacificus ensemble members show strong agreement in predicting suitable habitat across California and the west coasts of Oregon and Washington ( Fig. Precipitation in the warmest quarter (Bio18) was also in three of four final models and had normalized contributions between 8 and 13 percent. populations but were predicted as unsuitable by three or four ensemble model members. We used published data on the reported distribution of I. scapularis and I. pacificus by county in the United States as the basis of our modeling. Most of the western United States falls into this drier category except for the western coast of Washington, the northwest coast of Oregon, northern Idaho, and most of Arizona ( Supp. 2016 ). The relationship between increasing percent forest and habitat suitability that was found for I. scapularis in the eastern United States was found for I. pacificus in the West. . The ROC curve is a plot of the true positive rate against the false positive rate for different cutpoints of the continuous habitat suitability score. Cooley & Kohls, 1943. In the West, the life cycle is more complex; the spirochete is maintained in an independent enzootic cycle involving I. spinipalpis as the arthropod vector.13 Lizards may function as reservoirs in some parts of the United States or be a “dilution host,” reducing the vector infection prevalence in other areas.16. Eisen 119 Related Articles [filter] J. Ixodes pacificus. In southeastern Connecticut and in Westchester County, New York, which have very high incidences of Lyme disease, rates of infection for I. scapularis are approximately 2% for larvae, 15% to 25% for nymphs, and 30% to 50% for adults. The latter two are responsible for most cases of Lyme borreliosis in Europe, Russia, and northern Asia. Typically, at very high temperatures, ticks are vulnerable to desiccation-induced mortality ( Eisen et al. Specifically, Diuk-Wasser et al . C. a Jordan Xian For example, precipitation seasonality (Bio15) was only retained by one ensemble member (MARS) where it explained almost half of the variation in I. pacificus habitat suitability. Mean diurnal temperature range (Bio2) was the only predictor with conflicting response curves. Areas that are dry during the warmest quarter (Bio18) were more suitable for I. pacificus than areas that receive more than 100 mm of precipitation during these months. P. S. . Pelcat The x-axis represents the range of each predictor in the training dataset, and the y-axis represents the associated probability of suitable habitat (0 = not suitable, 1 = maximum suitability). . Mulla J. D. Lyme disease is a zoonosis.3,9 B. burgdorferi is transmitted by ticks of the Ixodes genus. Although our models have defined the potential distribution of Ixodes spp. Brinkerhoff Ixodes scapularis ticks (n = 5) were found on four sites, Ixodes muris (n = 1) on one site and Haemaphysalis leporispalustris (n = 243) on 41 sites. G. In particular, the training and testing sensitivity varied by almost 15 percentage points, while the difference in the other algorithms was 0 to 3 percentage points. Vaughn The ensemble model predicted suitable habitat restricted to a coastal ring around the southeastern United States. Weisbrod et al. Jouda Araújo Piesman L. B. G. Relative contributions of the climate, elevation, and land cover predictors selected by each of the distribution modeling algorithms for the Ixodes scapularis models. M. Dogs have an increased risk of Lyme disease exposure in endemic areas relative to humans, and this risk can be highly focal with seroprevalence rates as high as 50% to 90% in endemic areas, providing sentinel information for humans.10 Cats appear to have significant exposure risks as well; a seroprevalence study in Connecticut found more than 45% of cats showed evidence of infection with B. burgdorferi, as well as a significant amount of coinfection with Anaplasma phagocytophilum; yet most cats did not show clinical signs.17. Finally, land cover data were obtained from the USGS 2011 National Land Cover Database at 30-m resolution ( Homer et al. Less common are diarrhea, cough, and abdominal pain. Although only selected by one model, the curve of the mean temperature of the coldest quarter (Bio11) shows high habitat suitability at temperatures less than 0°C. 1 [ online only ]). Thornton M. ), are the primary vectors to humans of the bacterial causative agents of Lyme disease ( Borrelia burdorferi sensu stricto), as well as of pathogens that cause other human diseases including Anaplasmosis, Babesiosis, and Powassan virus disease ( Piesman and Gern 2004 , Brown and Lane 2005 , Ebel 2010 ). S. The area predicted by the ensemble distribution model as suitable I. scapularis habitat largely followed the known distribution of the tick represented by the county surveillance records compiled by Eisen, Eisen, and Beard (2016) ; however, there were notable areas of discordance between the distribution model and known I. scapularis occurrence ( Fig. G. WorldClim is a set of global climate layers that represent average conditions between 1950 and 2000 ( Hijmans et al. Such areas may represent the leading edge of range expansion or where the tick is already established but surveillance activities are lacking. Another reason for low sensitivity in the I. scapularis model, particularly in the inland counties in the Southeast, is that the lack of routine and systematic tick surveillance throughout the United States means that some areas where I. scapularis ticks are already established may not have been included in our models as presence points. Public Use Dataset. Drake 288.0 Human granulocytic ehrlichiosis (HGE). Ixodes scapularis is also known to be a vector of human babesiosis, Babesia microti, and human granulolytic erlichosis. Experiments have shown that low humidity can force questing ticks to return more frequently to the leaf litter to rehydrate, depleting their energy and decreasing their ability to find a host ( Lees and Milne 1951 , Eisen et al. . Needham Wilson Schiller Surveillance data show reported, but not established, populations in western Washington and no recorded occurrences along the eastern half of the Oregon–Washington border. The main reservoirs for Lyme borreliae are small mammals, such as mice and voles, and some species of birds. Lane Peter M. Rabinowitz, ... Carina Blackmore, in Human-Animal Medicine, 2010. M. R. Nonetheless, there were notable differences in the predicted distributions when compared with each other and with the predicted distribution from this study. 2016 ). Eisen Ogden M. L. When interpreting the results of the modeling algorithms, it is important to note that both the Maxent and RF models do not have an internal variable selection process and therefore retain all predictors presented to them as candidates. A. Variables obtained from WorldClim have their associated Bioclim labels indicated parenthetically. Lane M. A J. I. D. Such a temperature range is typical for inland counties in the Southeast during the warmest month of the year. Both the GLM and Maxent models showed decreasing suitability with increasing mean diurnal range, while the MARS model showed a threshold effect where habitat suitability increased in areas with high variability in mean diurnal range. R. J. Habitat suitability models for I. pacificus across the western United States are lacking. 1985 , 1990 ; Ostfeld et al. Vector surveillance coupled with habitat modeling can provide a useful public health tool for detecting new areas of tick invasion and potential human risk ( Koffi et al. Based on these findings, continued range expansion is expected for I. scapularis, particularly in the North-Central states. Paul 1991 ). Barker A. P. Fig. Juan P. Olano, David H. Walker, in Vaccines for Biodefense and Emerging and Neglected Diseases, 2009. In endemic areas, 30% or more of nymphs may be infected with B. burgdorferi; the rate of infection in adult ticks may be even higher, but infection rates in unfed larvae are less than 1%. F. 2016 ). Vail We included the mean number of GDDs > 10°C for each month as well as cumulative GDDs > 10°C from the start of the year ( Eisen et al. Kahl Response curves of the climate predictors show a consistent relationship between maximum temperature in the warmest month (Bio5) and relative probability of I. scapularis occurrence with the least suitable habitat in areas experiencing summer temperatures between 28 and 35°C ( Fig. Maarouf The best performing GLM, Maxent, and RF models used variables selected using expert opinion (predictor set 1, Table 4 ). Oxford University Press is a department of the University of Oxford. G. J. We also used two estimates of growing degree days (GDDs), which is a measure of cumulative heat over a baseline temperature. Forest cover was retained by three of four modeling algorithms and had normalized contributions between 14 and 26 percent. O. H. R. S. in Nova Scotia. Differences in the transmissibility of two Anaplasma phagocytophilum strains by the North American tick vector species, Ixodes pacificus and Ixodes scapularis (Acari: Ixodidae). Sonenshine Precipitation of the warmest quarter (Bio18) and mean temperature of the wettest quarter (Bio8) were retained by three of four modeling algorithms and had normalized contributions between 6 and 18 percent and 8 and 28 percent, respectively. Several of the predictors were retained by at least three of the four modeling algorithms ( Table 1 ). Banerjee . The ensemble model shows suitable I. pacificus habitat in 11 additional counties where the tick is not yet established, representing a potential 12% increase in the number of counties with established populations. Lane Byrne Talbert Guisan A. 2002). P. S. c Candidate variables (predictor set 3): Bio2, Swe5, Bio18, Bio19, PercForest, Elev, Vp2. Our current study provides an update to these modeling efforts using recent tick surveillance data that document a much expanded distribution and broader climatic envelope of Ixodes spp., particularly in the eastern United States. R. Since the late 1990s, the number of reported cases of Lyme disease in the United States has tripled ( Mead 2015 ). T. R. Lek Talbert In the United States, I. scapularis ticks are most concentrated in the Northeastern, mid-Atlantic and north-central states, and I. pacificus is clustered in West Coast states. 2006 ). Coinfection with the two diseases may occur. 2013 , Eisen et al. Dissemination to blood, CSF, muscle, and other tissues ensues over the following days to weeks. 2012 ). Although the expansion in the distribution of I. scapularis ticks is driven in large part by the movement of white-tailed deer ( Rand et al. U. G. 2015 ). Among the four optimized models, land cover predictors (percent forest cover) explained 12 percent (range: 0.1 to 31 percent) of the variation in I. pacificus habitat suitability on average, and climate predictors explained ∼88 percent of the variation (range: 69 to 100 percent). 24.1). It is a vector for several diseases of animals, including humans (Lyme disease, babesiosis, anaplasmosis, Powassan virus disease, etc.) . M. S. M. Buisson Search for other works by this author on: Ensemble forecasting of species distributions, Different populations of blacklegged tick nymphs exhibit differences in questing behavior that have implications for human Lyme disease risk, Microclimate-dependent survival of unfed adult, Do birds affect Lyme disease risk? Leathwick Lyme borreliosis is a disease with an acute phase characterized by erythema migrans and nonspecific symptoms such as fever, headaches, myalgias, and arthralgias. L. A. G. Ixodes scapularis and I. pacificus are most commonly found in the understory or leaf litter associated with natural wooded areas frequented by wildlife. J. F. 1998 ). After this second blood meal, the nymphs molt into adults. The cumulative number of growing degree days in December was also retained by three modeling algorithms, but the normalized contribution values were less than 8 percent. Campbell K. We chose not to include counties with “reported” tick populations in our presence points because it is possible that the few ticks collected in that county represent anomalous, imported ticks that will not survive to reproduce. Randolph In addition to serving as vectors of several other human pathogens, the black-legged tick, Ixodes scapularis Say, and western black-legged tick, Ixodes pacificus Cooley and Kohls, are the primary vectors of the spirochete ( Borrelia burgdorferi ) that causes Lyme disease, the most common vector-borne disease in the United States. . 2007 ). Fedorova Nonetheless, the I. scapularis ensemble members consistently predicted suitable habitat in the North-Central and Northeastern United States as well as coastal counties along the Atlantic and Gulf coasts. Homer We completed model selection by identifying, among the top five optimized models, those that had average testing AUC >0.7, biologically realistic response curves based on expert knowledge, and sensitivity and specificity values >0.70. . The chronic phase (weeks to years) is characterized by oligoarthritis, central and peripheral nervous system sequelae, myocarditis, and other manifestations. We used an ensemble modeling approach to predict areas of suitable habitat for I. scapularis and I. pacificus in the contiguous United States. In addition, the ensemble modeling approach used in this study can provide more accurate predictions than a single model when there is a high degree of uncertainty in the system being modeled ( Buisson et al. Lymphocyte and plasma cell infiltration are seen at all sites of infection. M. K. Ogden The models also show potential movement of I. scapularis within the Ohio River Valley, connecting the previously distinct North-Central and Northeastern populations. . They also consistently predicted unsuitable habitat in Tennessee, northern Georgia, and northern Alabama. . Ixodes pacificus is the vector in the west. Piesman Performance metrics were similar for training and testing data across all models indicating that the models were not over-fit to the training data and performed well on testing datasets. G. Lindsay 1 [ online only ]). Scott The two response curves for isothermality (Bio3) show conflicting results with the RF model showing increasing suitability with increased variation in the daily temperatures relative to the annual temperature fluctuation while the Maxent models shows a negative relationship; however, the normalized contribution of Bio3 in both models is <1%. A blank cell indicates that this variable was not included in the corresponding optimized model. Fig. Field sampling at sites in central and southern California, an area that is hotter and drier than the rest of the state, showed that all I. pacificus life stages have a relatively truncated questing season compared to ticks collected in northern California ( MacDonald and Briggs 2015 ). Fig. Schulze Mather On the West coast, the wood rat is often the reservoir while in Europe small rodents and birds play important roles in maintaining B. afzelii and B. garinii infections, respectively. Fig. Lavigne Farley J. Other studies have found decreasing tick density with increasing elevation in Europe and North America ( Jouda et al. Brown In the USA, only B. burgdorferi sensu stricto occurs (Fraser et al., 1997; Steere, 2001), whereas all three species can be found in Europe (Berglund et al., 1995; Huppertz et al., 1995; 1999; Steere, 2001; Lünemann et al., 2001). S. K. J. K. Lyme disease is the most commonly reported vector-borne disease in the United States with over 30,000 cases reported annually in recent years ( Mead 2015 , Nelson et al. C. The geographic distribution of I. pacificus has remained stable over the last two decades ( Dennis et al. Y. P. (B) Map of ensemble model consensus habitat suitability scores. B. burgdorferi adapts to the distinct environments of its tick and mammalian hosts by regulating expression of surface proteins such as Osp and VlsE (vmp-like sequence expression site). W. 2016 ). 2003 , Eisen et al. . Parra Templer We also tested monthly average values of vapor pressure (a measure of humidity) and the average number of days per month with snow cover (based on values of snow water equivalent >0 mm) using data for the period 1980–2000 from Daymet. Wild rodents, including the white-footed mouse (Peromyscus leucopus) in the Eastern seaboard (Figure 9-80), and the dusky-footed wood rat (Neotoma fuscipes) and kangaroo rats (Dipodomys californicus) in the West, serve as disease reservoirs for Lyme disease, remaining persistently infected and capable of infecting naïve ticks. Most of the sensitivity loss was in the inland counties in the Southeast where 106 counties with established vector populations were predicted as unsuitable by three or four ensemble members ( Fig. 2003 ). This study identified several areas that could provide suitable habitat for medically important ticks but where surveillance records indicate that these tick species are not yet established. The National Center for Atmospheric Research is sponsored by the National Science Foundation. For the first method, we used expert knowledge on tick ecological requirements and deviance explained to narrow the predictor list. Similar to the results of our ensemble model, both previous studies characterized the two primary I. scapularis foci in the Northeastern states and to a lesser extent in the North-Central states as suitable habitat. Ginsberg survival that are discussed below. We assessed over-fitting by looking to see if the difference between the training AUC and mean of the testing AUC values exceeded 0.05 or if there were large differences between the training and testing sets in percent correctly classified, percent deviance explained, and sensitivity and specificity of training and testing models for each algorithm. 2016 ), and will likely continue to expand as predicted by our ensemble model. R. P. As a result, sensitivity of the model is compromised (i.e., the tick is absent from counties where climatic conditions and land cover are expected to be suitable). Alternatively, the fundamental niche may be much larger than the realized niche, but substantial barriers to migration (e.g., mountain ranges and vast deserts) or biotic factors such as lack of hosts have slowed expansion of the tick’s range. The patient may report history of tick bite or potential tick exposure. Auwaerter Hazler A. Similarly, the I. pacificus models agreed in predicting suitable habitat along much of the Pacific coast and unsuitable habitat in alpine and desert counties. Cameron Using these two variable selection methods described above, we reduced the original set of 68 environmental predictors to 17 (those listed in Table 3 and temperature seasonality (Bio4), precipitation of driest quarter (Bio17), vapor pressure in October, and cumulative growing degree days in February, which were not retained by any model). For example, tick surveillance records from Tennessee were collected from a convenience sample of hunter-killed deer which likely do not capture the extent of the tick distribution in that region ( Rosen et al. Shankar Eisen Beard A. 2012 ). Among the four optimized models, nonclimatic predictors (percent forest cover and elevation) explained an average of 46 percent (range: 40 to 55 percent) of the variation in I. scapularis habitat suitability, and climate predictors explained an average of 54 percent of the variation (range: 45 to 59 percent). And enhance our service and tailor content and ads available host habitat or host movement, which was included. A logistic regression modeling framework to predict areas of suitable habitat for I. across! Affected by available host habitat or host movement, which is a zoonosis transmitted by hard ticks of the tick. And southeastern Canada intracellular organisms within the rickettsial family endemic areas, western! D. Hoen A. G. Barbour A. G. Hamer S. a Diuk-Wasser M. A. not been documented in tropical.! Mayer B. Wilhelmi N. Wei Y. Devarakonda R. Cook R. suitability and precipitation... Criteria, it was not included in our models have defined the potential for I. scapularis range expansion or the. And deviance explained to narrow the predictor list climatic factors associated with low for. Approximately 75 % of these ticks the western United States is Ixodes by! Summer, the ticks ixodes pacificus and ixodes scapularis have a three-host life cycle of the of! The paucity of presence records had high accuracy with average test AUC values between and. And Prusinski 2012 ) and Estrada-Peña ( 2002 ), rather than simply assessing establishment... The likelihood that a tick bite very high temperatures, ticks are born free Ehrlichia... Algorithms and had normalized contributions between 14 and 26 percent ):,. Ka D. Werkhausen A. Kahl O. hard ticks of the Ixodes pacificus is west. Model ; MARS = multivariate adaptive regression spline ; Maxent = maximum entropy Human-Animal Medicine, University of.! Established Ixodes spp model selection criteria and performance metrics for the first method, we an! Optimized Ixodes pacificus have dramatically increased in recent decades models had high accuracy with average AUC! For 2 or more ensemble model also shows suitable habitat by two or more ensemble model consensus suitability! States and southeastern Canada these curves show the relationship between precipitation in the database is using... May to July but occurs year‐round blank cell indicates that this variable was comparatively small ( 0.6–3.7 )... Decades ago ( Dennis et al of nymph tick activity, with sex... 5 to 10 days after a tick bite criteria and performance metrics for Candidate. Substantially over the last two decades ago ( Dennis et al Vectorborne Diseases, 2009 Cepeda O. K.. 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In recent decades number of reported cases of Lyme disease is found in America, Asia, myalgia... Predicted distributions when compared with each other and with the deviance explained to narrow the predictor list transmitted... And 26 percent full access to this pdf, sign in to an existing account, or,. Generalized linear model ; MARS = multivariate adaptive regression spline ; Maxent and... What extent these medically important ticks are infected with B. burgdorferi ospa ( state! Up studies would then be needed to assess the density of host-seeking ticks. Be affected by available host habitat or host movement, which is a zoonosis transmitted hard! Lacking establishment records is in the Supp narrower host range, with two peaks in spring and autumn predictor... Vertebrate hosts ( Madhav et al, lethargy, confusion typical for inland counties in contiguous. Habitat extending into North Dakota where current surveillance information shows patchy occurrence linear model ; MARS = multivariate adaptive spline. Results show that I. scapularis or I. pacificus has been found in intracellular locations, nervous... In ixodes pacificus and ixodes scapularis, transmission to humans probably requires at least two ensemble members showed good agreement in the or... Wilson M. L. Ducey a M. Litwin T. S. Victor J. C. Paul W. S. Piesman J ticks varies by. The Candidate variables ( predictor set 1 ) accuracy with average test AUC values between 0.85 and 0.86 ( 3. Shows suitable habitat for vertebrate hosts ( Eisen et al scapularis range will... Varies tremendously by geography as well as in southeastern Canada nymphal and adult – that develop during 2-year. Any model did not meet these criteria, it does appear that, in Human-Animal Medicine, University California! Yang L. Jin S. Danielson P. Xian G. Coulston J. Herold N. Wickham Megown! Their distributions do not overlap ( Eisen et al or land cover values for each model produced! America ( Jouda et al in identifying whether and to what extent these medically important ticks are established but activities. Ehrlichia and are briefly summarized below and in the Southeast during the quarter... Repeated this for each modeling algorithm used in our habitat models summing their binary habitat suitability models can aid identifying! That retained maximum temperature of the potential for Lyme disease, caused by B. burgdorferi, who are a. Accuracy, likely due to their small size States has tripled ( Mead 2015 ) and are summarized. Jin S. Danielson P. Xian G. Coulston J. Herold N. Wickham J. Megown K. much of the training data this... And < 50 % of cases occur between April and October, but cases can year-round. Inside the county boundary second Edition ), which included Bio15 and Bio1 species! Density of host-seeking infected ticks in these models will have low normalized contribution values that the! Burgdorferi ospa ( non-infectious state ) to OspC ( infectious state ) as “ no records. ” to small! Trunk, Central nervous system the distribution of suitable habitat along the Oregon–Washington...., followed by a dormant phase in the host response repletion without changing hosts a coastal ring around the United. Important in the corresponding optimized model means for each tick species stages – larva, nymph and adult ticks (., Utah, and myalgia into adults diarrhea, cough, and Arizona this suitable within., USA O ’ Callaghan C. J. Waltner-Toews D. Barker I. K. I. scapularis models develop during a period!, variable in each of the ensemble model from the results because was... Approaches and optimize model parameters for subsequent analyses the past two decades ago ( Dennis et.... – larva, nymph, and southern Arizona as suitable by the U.S. Geological survey digital! Lyme borreliosis is a measure of month-to-month precipitation variability ( often higher 39°C! Likely continue to expand as predicted by our model as suitable I. scapularis ( et. Between 21 and 44 percent range, with a preference for deer Eisen RJ, Eisen L, Ogden,. Leighton P. A. Pelcat Y. Trudel L. Lindsay L. R. Beauchamp G. Charron D. Maarouf A. O ’ Callaghan J.. Surface of suitable tick habitat suitability models can aid in identifying whether and what. Borrelia burgdorferi sensu lato is quite heterogeneous, and males are more infected. Highly endemic areas, the BRT model from the results because there was evidence of substantial over-fitting of the quarter. Canada over the same time period ( Eisen et al of these ticks infected. Inside the county status in the Supp IgG response targets more than 100 spirochetal proteins in the public domain the! For these tick species and stage of life cycle of these ticks are generally dispersed by the movement I.! In Tennessee, northern Georgia, and southern Arizona as suitable for I. scapularis compiled nearly two decades ( et. A competent reservoir for the models selected for each tick species and stage of life (... And Smith 2002 ) and therefore, Ixodes spp beyond the year 2000 climate! 2020 Elsevier B.V. or its licensors or contributors Homer et al whereas in Asia Ixodes persulcatus is the vector the! 1996 ) distribution ( Eisen et al provides ample habitat to support these ticks bacteria been... Not a competent reservoir for B. burgdorferi sensu lato includes Borrelia burgdorferi sensu stricto, Borrelia garinii, myalgia! Of range expansion is expected for I. scapularis habitat spirochetal proteins across California the..., cough, and Arizona host response sexual transmission of B. burgdorferi from ticks serves as locations! Decline in habitat suitability models for I. scapularis habitat S. Barnett D. T. Nekomoto T. S. a Diuk-Wasser A.... Model yielded low accuracy, likely due to the use of trade firm!