Geology topics

Jean I. Tsao

Publications and source records attributed to Jean I. Tsao.

11 recordsLinked to original sources

USGS2021

Selective host attachment by Ixodes scapularis (Acari: Ixodidae): Tick-lizard associations in the southeastern United States

Questing behavior and host associations of immature blacklegged ticks, Ixodes scapularis Say, from the southeastern United States are known to differ from those in the north. To elucidate these relationships we describe host associations of larval and nymphal I. scapularis from 8 lizard species sampled from 5 sites in the southeastern U.S. Larvae and nymphs attached in greater numbers to larger lizards than to smaller lizards, with differential levels of attachment to different lizard species. Blacklegged ticks are generally attached to skinks of the genus Plestiodon in greater numbers per unit lizard weight than to anoles ( Anolis ) or fence lizards ( Sceloporus ). The broad-headed skink, Plestiodon laticeps (Schneider), was a particularly important host for immature I. scapularis in our study and in several previous studies of tick–host associations in the southeast. Blacklegged ticks show selective attachment to Plestiodon lizard hosts in the southeast, but whether this results from behavioral host preferences or from ecological factors such as timing or microhabitat distributions of tick questing and host activity remains to be determined.

Alabama, Florida, North Carolina, South Carolina, ↗

USGS2021

Why Lyme disease is common in the northern US, but rare in the south: The roles of host choice, host-seeking behavior, and tick density

Lyme disease is common in the northeastern United States, but rare in the southeast, even though the tick vector is found in both regions. Infection prevalence of Lyme spirochetes in host-seeking ticks, an important component to the risk of Lyme disease, is also high in the northeast and northern midwest, but declines sharply in the south. As ticks must acquire Lyme spirochetes from infected vertebrate hosts, the role of wildlife species composition on Lyme disease risk has been a topic of lively academic discussion. We compared tick–vertebrate host interactions using standardized sampling methods among 8 sites scattered throughout the eastern US. Geographical trends in diversity of tick hosts are gradual and do not match the sharp decline in prevalence at southern sites, but tick–host associations show a clear shift from mammals in the north to reptiles in the south. Tick infection prevalence declines north to south largely because of high tick infestation of efficient spirochete reservoir hosts (rodents and shrews) in the north but not in the south. Minimal infestation of small mammals in the south results from strong selective attachment to lizards such as skinks (which are inefficient reservoirs for Lyme spirochetes) in the southern states. Selective host choice, along with latitudinal differences in tick host-seeking behavior and variations in tick densities, explains the geographic pattern of Lyme disease in the eastern US.

PLoS Biology↗

USGS2021

Seasonality of acarological risk of exposure to Borrelia miyamotoi from questing life stages of Ixodes scapularis collected from Wisconsin and Massachusetts, USA

Measures of acarological risk of exposure to Ixodes scapularis -borne disease agents typically focus on nymphs; however, the relapsing fever group spirochete, Borrelia miyamotoi can be transmitted transovarially, and I. scapularis larvae are capable of transmitting B. miyamotoi to their hosts. To quantify the larval contribution to acarological risk, relative to nymphs and adults, we collected questing I. scapularis for 3 yr at Fort McCoy, Wisconsin (WI, n = 23,367 ticks), and Cape Cod, Massachusetts (MA, n = 4,190) in the United States. Borrelia miyamotoi infection prevalence was estimated for I. scapularis larvae, nymphs, females, and males, respectively, as 0.88, 2.05, 0.63, and 1.22% from the WI site and 0.33, 2.32, 2.83, and 2.11% from the MA site. Densities of B. miyamotoi -infected ticks (DIT, per 1,000 m 2 ) were estimated for larvae, nymphs, females, and males, respectively, as 0.36, 0.14, 0.01, and 0.03 from the WI site and 0.05, 0.06, 0.03, and 0.02 from the MA site. Thus, although larval infection prevalence with B. miyamotoi was significantly lower than that of nymphs and similar to that of adults, because of their higher abundance, the larval contribution to the overall DIT was similar to that of nymphs and trended towards a greater contribution than adults. Assuming homogenous contact rates with humans, these results suggest that eco-epidemiological investigations of B. miyamotoi disease in North America should include larvae.A fuller appreciation of the epidemiological implications of these results, therefore, requires an examination of the heterogeneity in contact rates with humans among life stages.

Massachusetts, Wisconsin↗

USGS2020

Possible effects of climate change on ixodid ticks and the pathogens they transmit: Predictions and observations

The global climate has been changing over the last century due to greenhouse gas emissions and will continue to change over this century, accelerating without effective global efforts to reduce emissions. Ticks and tick-borne diseases (TTBDs) are inherently climate-sensitive due to the sensitivity of tick lifecycles to climate. Key direct climate and weather sensitivities include survival of individual ticks, and the duration of development and host-seeking activity of ticks. These sensitivities mean that in some regions a warming climate may increase tick survival, shorten life-cycles and lengthen the duration of tick activity seasons. Indirect effects of climate change on host communities may, with changes in tick abundance, facilitate enhanced transmission of tick-borne pathogens. High temperatures, and extreme weather events (heat, cold, and flooding) are anticipated with climate change, and these may reduce tick survival and pathogen transmission in some locations. Studies of the possible effects of climate change on TTBDs to date generally project poleward range expansion of geographical ranges (with possible contraction of ranges away from the increasingly hot tropics), upslope elevational range spread in mountainous regions, and increased abundance of ticks in many current endemic regions. However, relatively few studies, using long-term (multi-decade) observations, provide evidence of recent range changes of tick populations that could be attributed to recent climate change. Further integrated ‘One Health’ observational and modeling studies are needed to detect changes in TTBD occurrence, attribute them to climate change, and to develop predictive models of public- and animal-health needs to plan for TTBD emergence.

Journal of Medical Entomology↗

USGS2020

Local abundance of Ixodes scapularis in forests: Effects of environmental moisture, vegetation characteristics, and host abundance

Ixodes scapularis is the primary vector of Lyme disease spirochetes in eastern and central North America, and local densities of this tick can affect human disease risk. We sampled larvae and nymphs from sites in Massachusetts and Wisconsin, USA, using flag/drag devices and by collecting ticks from hosts, and measured environmental variables to evaluate the environmental factors that affect local distribution and abundance of I. scapularis . Our sites were all forested areas with known I. scapularis populations. Environmental variables included those associated with weather (e.g., temperature and relative humidity), vegetation characteristics (at canopy, shrub, and ground levels), and host abundance (small and medium-sized mammals and reptiles). The numbers of larvae on animals at a given site and season showed a logarithmic relationship to the numbers in flag/drag samples, suggesting limitation in the numbers on host animals. The numbers of nymphs on animals showed no relationship to the numbers in flag/drag samples. These results suggest that only a small proportion of larvae and nymphs found hosts because in neither stage did the numbers of host-seeking ticks decline with increased numbers on hosts. Canopy cover was predictive of larval and nymphal numbers in flag/drag samples, but not of numbers on hosts. Numbers of small and medium-sized mammal hosts the previous year were generally not predictive of the current year’s tick numbers, except that mouse abundance predicted log numbers of nymphs on all hosts the following year. Some measures of larval abundance were predictive of nymphal numbers the following year. The mean number of larvae per mouse was well predicted by measures of overall larval abundance (based on flag/drag samples and samples from all hosts), and some environmental factors contributed significantly to the model. In contrast, the mean numbers of nymphs per mouse were not well predicted by environmental variables, only by overall nymphal abundance on hosts. Therefore, larvae respond differently than nymphs to environmental factors. Furthermore, flag/drag samples provide different information about nymphal numbers than do samples from hosts. Flag/drag samples can provide information about human risk of acquiring nymph-borne pathogens because they provide information on the densities of ticks that might encounter humans, but to understand the epizootiology of tick-borne agents both flag/drag and host infestation data are needed.

Massachusetts, Wisconsin↗

USGS2018

Evidence for geographic variation in life-cycle processes affecting phenology of the Lyme disease vector Ixodes scapularis (Acari: Ixodidae) in the United States

The seasonal activity pattern of immature Ixodes scapularis Say (Acari: Ixodidae) varies geographically in the United States, which may affect the efficiency of transmission cycles of pathogens transmitted by this species. To study the factors that determine seasonality, a multiyear study at seven sites across the geographic range of I. scapularis systematically collected questing ticks by flagging/dragging, and feeding ticks by capture of their hosts. The observed phenology patterns were consistent with previous studies reporting geographic variation in seasonal tick activity. Predictions of seasonal activity for each site were obtained from an I. scapularis simulation model calibrated using contemporaneous weather data. A range of scenarios for life-cycle processes—including different regimes of temperature-independent behavioral and developmental diapause, variations in temperature–development rate relationships, and temperature-dependent tick activity—were used in model formulations. These formulations produced a range of simulations of seasonal activity for each site and were compared against the field observed tick data using negative binomial regression models. Best fit scenarios were chosen for each site on the basis of Akaike’s information criterion and regression model parameters. This analysis suggests that temperature-independent diapause mechanisms explain some key observed variations in I. scapularis seasonality, and are responsible in part for geographic variations in I. scapularis seasonality in the United States. However, diapause appears to operate in idiosyncratic ways in different regions of the United States, so further studies on populations in different regions will be needed to enable predictive modeling of climatic and climate change effects on I. scapularis seasonal activity and pathogen transmission.

Journal of Medical Entomology↗

USGS2017

Environmental factors affecting survival of immature Ixodes scapularis and implications for geographical distribution of lyme disease: The climate/behavior hypothesis

Recent reports suggest that host-seeking nymphs in southern populations of Ixodes scapularis remain below the leaf litter surface, while northern nymphs seek hosts on leaves and twigs above the litter surface. This behavioral difference potentially results in decreased tick contact with humans in the south, and fewer cases of Lyme disease. We studied whether north-south differences in tick survival patterns might contribute to this phenomenon. Four month old larvae resulting from a cross between Wisconsin males and South Carolina females died faster under southern than under northern conditions in the lab, as has previously been reported for ticks from both northern and southern populations. However, newly-emerged larvae from Rhode Island parents did not differ consistently in mortality under northern and southern conditions, possibly because of their younger age. Survival is lower, and so the north-south survival difference might be greater in older ticks. Larval survival was positively related to larval size (as measured by scutal area), while survival was positively related to larval fat content in some, but not all, trials. The difference in larval survival under northern vs. southern conditions might simply result from faster metabolism under warmer southern conditions leading to shorter life spans. However, ticks consistently died faster under southern than under northern conditions in the laboratory when relative humidity was low (75%), but not under moderate (85%) or high (95%) RH. Therefore, mortality due to desiccation stress is greater under southern than under northern conditions. We hypothesize that mortality resulting from the greater desiccation stress under southern conditions acts as a selective pressure resulting in the evolution of host-seeking behavior in which immatures remain below the leaf litter surface in southern I . scapularis populations, so as to avoid the desiccating conditions at the surface. If this hypothesis is correct, it has implications for the effect of climate change on the future distribution of Lyme disease.

PLoS ONE↗

USGS2015

Different populations of blacklegged tick nymphs exhibit differences in questing behavior that have implications for human lyme disease risk

Animal behavior can have profound effects on pathogen transmission and disease incidence. We studied the questing (= host-seeking) behavior of blacklegged tick ( Ixodes scapularis ) nymphs, which are the primary vectors of Lyme disease in the eastern United States. Lyme disease is common in northern but not in southern regions, and prior ecological studies have found that standard methods used to collect host-seeking nymphs in northern regions are unsuccessful in the south. This led us to hypothesize that there are behavior differences between northern and southern nymphs that alter how readily they are collected, and how likely they are to transmit the etiological agent of Lyme disease to humans. To examine this question, we compared the questing behavior of I. scapularis nymphs originating from one northern (Lyme disease endemic) and two southern (non-endemic) US regions at field sites in Wisconsin, Rhode Island, Tennessee, and Florida. Laboratory-raised uninfected nymphs were monitored in circular 0.2 m 2 arenas containing wooden dowels (mimicking stems of understory vegetation) for 10 (2011) and 19 (2012) weeks. The probability of observing nymphs questing on these stems (2011), and on stems, on top of leaf litter, and on arena walls (2012) was much greater for northern than for southern origin ticks in both years and at all field sites (19.5 times greater in 2011; 3.6-11.6 times greater in 2012). Our findings suggest that southern origin I. scapularis nymphs rarely emerge from the leaf litter, and consequently are unlikely to contact passing humans. We propose that this difference in questing behavior accounts for observed geographic differences in the efficacy of the standard sampling techniques used to collect questing nymphs. These findings also support our hypothesis that very low Lyme disease incidence in southern states is, in part, a consequence of the type of host-seeking behavior exhibited by southern populations of the key Lyme disease vector.

PLoS ONE↗

USGS2014

Comparison of survival patterns of northern and southern genotypes of the North American tick Ixodes scapularis (Acari: Ixodidae) under northern and southern conditions

Background Several investigators have reported genetic differences between northern and southern populations of Ixodes scapularis in North America, as well as differences in patterns of disease transmission. Ecological and behavioral correlates of these genetic differences, which might have implications for disease transmission, have not been reported. We compared survival of northern with that of southern genotypes under both northern and southern environmental conditions in laboratory trials. Methods Subadult I. scapularis from laboratory colonies that originated from adults collected from deer from several sites in the northeastern, north central, and southern U.S. were exposed to controlled conditions in environmental chambers. Northern and southern genotypes were exposed to light:dark and temperature conditions of northern and southern sites with controlled relative humidities, and mortality through time was recorded. Results Ticks from different geographical locations differed in survival patterns, with larvae from Wisconsin surviving longer than larvae from Massachusetts, South Carolina or Georgia, when held under the same conditions. In another experiment, larvae from Florida survived longer than larvae from Michigan. Therefore, survival patterns of regional genotypes did not follow a simple north–south gradient. The most consistent result was that larvae from all locations generally survived longer under northern conditions than under southern conditions. Conclusions Our results suggest that conditions in southern North America are less hospitable than in the north to populations of I. scapularis . Southern conditions might have resulted in ecological or behavioral adaptations that contribute to the relative rarity of I. scapularis borne diseases, such as Lyme borreliosis, in the southern compared to the northern United States.

Parasites & Vectors↗

USGS2014

Minimal role of eastern fence lizards in Borrelia burgdorferi transmission in central New Jersey oak/pine woodlands

The Eastern fence lizard, Sceloporus undulatus , is widely distributed in eastern and central North America, ranging through areas with high levels of Lyme disease, as well as areas where Lyme disease is rare or absent. We studied the potential role of S. undulatus in transmission dynamics of Lyme spirochetes by sampling ticks from a variety of natural hosts at field sites in central New Jersey, and by testing the reservoir competence of S. undulatus for Borrelia burgdorferi in the laboratory. The infestation rate of ticks on fence lizards was extremely low (proportion infested = 0.087, n = 23) compared to that on white footed mice and other small mammals (proportion infested = 0.53, n = 140). Of 159 nymphs that had fed as larvae on lizards that had previously been exposed to infected nymphs, none was infected with B. burgdorferi , compared with 79.9% of 209 nymphs that had fed as larvae on infected control mice. Simulations suggest that changes in the numbers of fence lizards in a natural habitat would have little effect on the infection rate of nymphal ticks with Lyme spirochetes. We conclude that in central New Jersey S. undulatus plays a minimal role in the enzootic transmission cycle of Lyme spirochetes.

New Jersey↗

USGS2013

Flagging versus dragging as sampling methods for nymphal Ixodes scapularis (Acari: Ixodidae)

The nymphal stage of the blacklegged tick, Ixodes scapularis (Acari: Ixodidae), is responsible for most transmission of Borrelia burgdorferi, the etiologic agent of Lyme disease, to humans in North America. From 2010 to fall of 2012, we compared two commonly used techniques, flagging and dragging, as sampling methods for nymphal I. scapularis at three sites, each with multiple sampling arrays (grids), in the eastern and central United States. Flagging and dragging collected comparable numbers of nymphs, with no consistent differences between methods. Dragging collected more nymphs than flagging in some samples, but these differences were not consistent among sites or sampling years. The ratio of nymphs collected by flagging vs dragging was not significantly related to shrub density, so habitat type did not have a strong effect on the relative efficacy of these methods. Therefore, although dragging collected more ticks in a few cases, the numbers collected by each method were so variable that neither technique had a clear advantage for sampling nymphal I. scapularis.

Journal of Vector Ecology↗