[Show abstract][Hide abstract] ABSTRACT: 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 m2 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 accountsfor 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 hostseeking behavior exhibited by southern populations of the key Lyme disease vector.
PLoS ONE 05/2015; 10(5):e0127450. DOI:10.1371/journal.pone.0127450 · 3.23 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: The European sheet-web spider Linyphia triangularis (Araneae: Linyphiidae) has become established in Maine, where it often reaches very high densities. Two lines of evidence from previous work suggest that L. triangularis affects populations of the native linyphiid spider Frontinella communis. First, F. communis individuals are relatively scarce in both forest and coastal habitat where L. triangularis is common, but more common where L. triangularis is at low density. Second, in field experiments, F. communis species are less likely to settle in experimental plots when L. triangularis is present, and F. communis disappears from study plots when L. triangularis is introduced. Here we test two mechanisms that may underlie these patterns. First, we tested whether L. triangularis invades and usurps the webs of F. communis. When spiders were released onto webs of heterospecifics, L. triangularis was more likely to take over or share webs of F. communis than the reverse. We also observed natural takeovers of F. communis webs. Second, we explored the hypothesis that L. triangularis reduces prey availability for native species. We sampled flying prey in areas with L. triangularis and those where it had been removed, and found no effect of spider presence on measured prey density. We also found no effect of prey supplementation on web tenacity in F. communis, suggesting that F. communis movements are not highly dependent on prey availability. We conclude that web takeover is likely more important than prey reduction in driving negative effects of L. triangularis on F. communis.
[Show abstract][Hide abstract] ABSTRACT: Ambient temperature can influence tick development time, and can potentially affect tick interactions with pathogens and with vertebrate hosts. We studied the effect of ambient temperature on duration of attachment of larval blacklegged ticks, Ixodes scapularis Say, to eastern fence lizards, Sceloporus undulatus (Bosc & Daudin). Feeding periods of larvae that attached to lizards under preferred temperature conditions for the lizards (WARM treatment: temperatures averaged 36.6°C at the top of the cage and 25.8°C at the bottom, allowing behavioral thermoregulation) were shorter than for larvae on lizards held under cool conditions (COOL treatment temperatures averaged 28.4°C at top of cage and 24.9°C at the bottom). The lizards were infested with larvae four times at roughly monthly intervals. Larval numbers successfully engorging and dropping declined and feeding period was longer after the first infestation.
Journal of Medical Entomology 11/2014; 51(6). DOI:10.1603/ME14068 · 1.82 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: 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.
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.
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.
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.
[Show abstract][Hide abstract] ABSTRACT: Laboratory studies have shown clear relationships between relative humidity (RH) and the activity and survival of Ixodes scapularis Say (blacklegged tick). However, field studies have produced conflicting results. We examined this relationship using weekly tick count totals and hourly RH observations at three field sites, stratified by latitude, within the state of Rhode Island. Records of nymphal tick abundance were compared with several RH-related variables (e.g., RH at time of sampling and mean weekly daytime RH). In total, 825 nymphs were sampled in 2009, a year of greater precipitation, with a weighted average leaf litter RH recorded at time of sampling of 85.22%. Alternatively, 649 nymphs were collected in 2010, a year of relatively low precipitation, and a weighted average RH recorded at time of sampling was 75.51%. Negative binomial regression analysis of tick count totals identified cumulative hours P = 0.0037; 2010: P P = 0.0016) in 2010. These results suggest a lag effect between moisture availability and patterns of tick activity and abundance. Differences in the relative importance of each RH variable between years may have been due to abnormally wet summer conditions in 2009.
Journal of Medical Entomology 07/2014; 51(4). DOI:10.1603/ME13186 · 1.82 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: Abstract 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.
Journal of Parasitology 05/2014; 100(5). DOI:10.1645/14-503.1 · 1.26 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: The numbers of human cases of Lyme disease and other tick-borne diseases (TBDs) reported each
year to CDC have been increasing steadily in the United States (US), currently totaling tens of thousands of
diagnosed human cases annually (Table 1). The reasons behind this increase are complex and involve
multiple factors including: (1) ecological changes, possibly including climate change, and shifts in land use
patterns, (2) increasing deer and wildlife populations and closer associations between humans and wildlife,
(3) human behavioral changes that have led to greater exposure risks, and (4) improvements in disease
diagnosis, surveillance, and reporting practices
[Show abstract][Hide abstract] ABSTRACT: Lyme borreliosis (LB) is the most commonly reported vector-borne disease in north temperate regions worldwide, affecting an estimated 300,000 people annually in the United States alone. The incidence of LB is correlated with human exposure to its vector, the blacklegged tick (Ixodes scapularis). To date, attempts to model tick encounter risk based on environmental parameters have been equivocal. Previous studies have not considered (1) the differences between relative humidity (RH) in leaf litter and at weather stations, (2) the RH threshold that affects nymphal blacklegged tick survival, and (3) the time required below the threshold to induce mortality. We clarify the association between environmental moisture and tick survival by presenting a significant relationship between the total number of tick adverse moisture events (TAMEs - calculated as microclimatic periods below a RH threshold) and tick abundance each year.
We used a 14-year continuous statewide tick surveillance database and corresponding weather data from Rhode Island (RI), USA, to assess the effects of TAMEs on nymphal populations of I. scapularis. These TAMEs were defined as extended periods of time (>8 h below 82% RH in leaf litter). We fit a sigmoid curve comparing weather station data to those collected by loggers placed in tick habitats to estimate RH experienced by nymphal ticks, and compiled the number of historical TAMEs during the 14-year record.
The total number of TAMEs in June of each year was negatively related to total seasonal nymphal tick densities, suggesting that sub-threshold humidity episodes >8 h in duration naturally lowered nymphal blacklegged tick abundance. Furthermore, TAMEs were positively related to the ratio of tick abundance early in the season when compared to late season, suggesting that lower than average tick abundance for a given year resulted from tick mortality and not from other factors.
Our results clarify the mechanism by which environmental moisture affects blacklegged tick populations, and offers the possibility to more accurately predict tick abundance and human LB incidence. We describe a method to forecast LB risk in endemic regions and identify the predictive role of microclimatic moisture conditions on tick encounter risk.
[Show abstract][Hide abstract] ABSTRACT: Background/Question/Methods
In the eastern US, most human cases occur in the North, despite the vector – the blacklegged tick- being found throughout the eastern U.S. Variation in the abundance of infected ticks also exists among Lyme disease endemic regions, as well where the blacklegged tick is actively expanding. A central question in Lyme disease ecology is how does biodiversity affect human disease risk?
To explain this pronounced North-South gradient in Lyme disease, medical entomologists conventionally hypothesize that nymphal ticks (the epidemiologically riskiest life stage) exhibit a different host-seeking behavior in the South such that the risk of humans being bitten by an infected nymph is negligible. An alternative hypothesis has been that juvenile ticks in the South disproportionately feed on incompetent hosts such as lizards, rather than on reservoir competent hosts; resulting in few infected nymphs. This hypothesis has been broadened into a more general biodiversity-dilution hypothesis, whereby increased host biodiversity is thought to reduce the density of infected ticks.
We used a modeling approach to explore the consequences of biodiversity on the relative abundance of infected nymphs. We modified a previously published model that investigates transmission between a seasonally dynamic reservoir host population and a seasonally dynamic vector tick population.
Our model shows that either reduction or amplification of the density of infected nymphs can occur with increased host biodiversity, depending on the biological details of the modeled host community. If competition exerted by alternative incompetent hosts reduces the density of reservoir hosts, dilution may occur. Similarly, if alternative hosts feed juvenile ticks at a much higher rate such that they divert juvenile ticks away from reservoir hosts, dilution may occur. Finally, if alternative hosts kill juvenile ticks that attempt to feed on them, for example, through grooming or through an immune response, dilution can also occur. If alterative hosts, however, allow a greater proportion of juvenile ticks to feed successfully, then amplification may occur.
In addition to these model results, empirical results for a multisite (n=9) study conducted to test the biodiversity- dilution hypothesis will be discussed in the context of the model as well as of other studies addressing the relationship between biodiversity and Lyme disease.
[Show abstract][Hide abstract] ABSTRACT: 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.
[Show abstract][Hide abstract] ABSTRACT: West Nile Virus (WNV) and Eastern Equine Encephalitis Virus (EEEV) are both primarily bird viruses, which can be transmitted by several mosquito species. Differences in larval habitats, flight, and biting patterns of the primary vector species result in substantial differences in epidemiology, with WNV more common, primarily occurring in urban areas, and EEEV relatively rare, typically occurring near swamp habitats. The complex transmission ecology of these viruses complicates prediction of disease outbreaks. The Rhode Island Department of Environmental Management (DEM) and Department of Health (DoH) provide prevention assistance to towns and maintain a mosquito surveillance program to identify potential disease risk. Responses to potential outbreaks follow a protocol based on surveillance results, assessment of human risk, and technical consultation.
Rhode Island medical journal 01/2013; 96(7):33-35.
[Show abstract][Hide abstract] ABSTRACT: Invasive spiders pose potential threats to native spiders. In 2002, the European spider Linyphia triangularis (Clerck, 1757) (Araneae: Linyphiidae) was discovered in all but one county in Maine. At Acadia National Park, we conducted a 4-year study of L. triangularis and three native linyphiid species of a similar size (Frontinella communis (Hentz, 1850), Pityohyphantes subarcticus Chamberlin and Ivie, 1943, and Neriene radiata (Walckenaer, 1842)). Using line-transect surveys, we measured population densities in coastal and forest habitat. The density of L. triangularis varied across years but was always significantly higher on the coast than in the forest. In contrast, only one native species was present on the coast and at very low numbers. Coastal L. triangularis were larger and in better condition than those in the forest, and numbers and biomass of insect prey were also higher on the coast. In 2 years, we also conducted transects at a second coastal location in Maine where the invader was at low density. At that site, native densities were substantially higher than at either Acadia site. Our data are consistent with the hypothesis that L. triangularis is reducing populations of native spiders. Companion studies suggest that L. triangularis negatively impacts natives by usurping both web sites and webs.
Canadian Journal of Zoology 07/2011; 89(8):668-677. DOI:10.1139/z11-050 · 1.35 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: The mosquito larvicide methoprene is a juvenile growth hormone mimic that is widely used to control mosquito larvae in stormwater catch basins. This study addresses two concerns pertaining to methoprene's use for mosquito control. First, measurements of methoprene concentrations were made from water in catch basins that had been treated with methoprene and from an adjoining salt pond near where the treated catch basins emptied. The concentrations of methoprene in catch basins and at drainage outlets after application at the rates currently used for mosquito control in southern Rhode Island were 0.5 ppb and lower, orders of magnitude below what has been determined as detrimental to organisms other than mosquitoes. Second, the effects of methoprene on the communities that live in catch basins were evaluated both in simulated catch basins in the laboratory and in actual catch basins in the field. We found no evidence of declines in abundances of any taxa attributable to the application. Furthermore, we found no consistent changes in community-level parameters (e.g., taxonomic richness, and dominance-diversity relationships) related to methoprene application in either field or laboratory trials.
[Show abstract][Hide abstract] ABSTRACT: Natal habitat use by dragonflies was assessed on an urban to rural land-use gradient at a set of 21 wetlands, during two emergence
seasons (2004, 2005). The wetlands were characterized for urbanization level by using the first factor from a principal components
analysis combining chloride concentration in the wetland and percent forest in the surrounding buffer zone. Measurements of
species diversity and its components (species richness and evenness) were analyzed and compared along the urbanization gradient,
as were distributions of individual species. Dragonfly diversity, species richness, and evenness did not change along the
urbanization gradient, so urban wetlands served as natal habitat for numerous dragonfly species. However, several individual
species displayed strong relationships to the degree of urbanization, and most were more commonly found at urban sites and
at sites with fish. In contrast, relatively rare species were generally found at the rural end of the gradient. These results
suggest that urban wetlands can play important roles as dragonfly habitat and in dragonfly conservation efforts, but that
conservation of rural wetlands is also important for some dragonfly species.
[Show abstract][Hide abstract] ABSTRACT: Acquisition of ticks by bird hosts is a central process in the transmission cycles of many tick-borne zoonoses, but tick recruitment by birds has received little direct study. We documented acquisition of Ixodes scapularis Say on birds at Fire Island, NY, by removing ticks from mist-netted birds, and recording the number of ticks on birds recaptured within 4 d of release. Eight bird species acquired at least 0.8 ticks bird(-1) day(-1) during the seasonal peak for at least one age class of I. scapularis. Gray Catbirds, Eastern Towhees, Common Yellowthroats, and Northern Waterthrushes collectively accounted for 83% of all tick acquisitions; and six individuals apportioned among Black-billed Cuckoo, Gray Catbird, Eastern Towhee, and Common Yellowthroat were simultaneously infested with both larvae and nymphs. Bird species with the highest acquisition rates were generally ground foragers, whereas birds that did not acquire ticks in our samples generally foraged above the ground. Tick acquisition by birds did not differ between deciduous and coniferous forests. Among the 15 bird species with the highest recruitment rates, acquisition of nymphs was not correlated with acquisition of larvae. Tick acquisition rates by individual bird species were not correlated with the reservoir competence of those species for Lyme borreliae. However, birds with high tick acquisition rates can contribute large numbers of infected ticks, and thus help maintain the enzootic cycle, even if their levels of reservoir competence are relatively low.
Journal of Medical Entomology 11/2010; 47(6):1019-27. DOI:10.1603/ME10086 · 1.82 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: Utility of early-season mosquito surveillance to predict West Nile virus activity in late summer was assessed in Suffolk County, NY. Dry ice-baited CDC miniature light traps paired with gravid traps were set weekly. Maximum-likelihood estimates of WNV positivity, minimum infection rates, and % positive pools were generally well correlated. However, positivity in gravid traps was not correlated with positivity in CDC light traps. The best early-season predictors of WNV activity in late summer (estimated using maximum-likelihood estimates of Culex positivity in August and September) were early date of first positive pool, low numbers of mosquitoes in July, and low numbers of mosquito species in July. These results suggest that early-season entomological samples can be used to predict WNV activity later in the summer, when most human cases are acquired. Additional research is needed to establish which surveillance variables are most predictive and to characterize the reliability of the predictions.
[Show abstract][Hide abstract] ABSTRACT: : Mosquito host-seeking activity was studied using a custom-designed trap to explore: (1) at which time interval of the night adult mosquito abatement would be most effective, and (2) if there exists an avian-specific host-seeking preference. Overnight trials using traps baited with dry ice showed that Aedes taeniorhynchus (Wiedemann) was most active at dusk and was then captured throughout the night. In contrast, Culex spp. (Cx. pipiens (Linnaeus) and Cx. restuans (Theobald) delayed most activity until about two h after dusk and were then captured through the night. This pattern suggests that management activities directed at adult Culex spp. would be most effective if initiated well after sunset. Mosquito capture rates in traps baited with birds in net bags were significantly greater than those with empty net bags, indicating that mosquitoes were attracted to the birds and not incidentally being sucked in by the custom trap's strong fan motor (Wilcoxon matched-pairs signed-ranks test, n= 24, t= 30, p < 0.05). Regression analysis showed that bird weight influenced mosquito attraction (r2= 0.21, p= 0.02). Trials with paired traps that contained different native bird species showed that Gray Catbirds, Dumatella carolinensis, attracted more mosquitoes than the heavier Northern Cardinals, Cardinalis cardinalis (paired samples t-test, t= 2.58, df= 7, p= 0.04). However, attractiveness did not differ substantially among bird species, and Gray Catbirds did not attract more mosquitoes than all other birds combined as a group. American Robins, Turdus migratorius (n = 4) were comparable in attractiveness to other bird species, but not enough American Robins were captured for a comprehensive study of mosquito avian preference.
[Show abstract][Hide abstract] ABSTRACT: Mosquito host-seeking activity was studied using a custom-designed trap to explore: (1) at which time interval of the night adult mosquito abatement would be most effective, and (2) if there exists an avian-specific host-seeking preference. Overnight trials using traps baited with dry ice showed that Aedes taeniorhynchus (Wiedemann) was most active at dusk and was then captured throughout the night. In contrast, Culex spp. (Cx. pipiens (Linnaeus) and Cx. restuans (Theobald) delayed most activity until about two h after dusk and were then captured through the night. This pattern suggests that management activities directed at adult Culex spp. would be most effective if initiated well after sunset. Mosquito capture rates in traps baited with birds in net bags were significantly greater than those with empty net bags, indicating that mosquitoes were attracted to the birds and not incidentally being sucked in by the custom trap's strong fan motor (Wilcoxon matched-pairs signed-ranks test, n=24, t=30, p<0.05). Regression analysis showed that bird weight influenced mosquito attraction (r2=0.21, p=0.02). Trials with paired traps that contained different native bird species showed that Gray Catbirds, Dumatella carolinensis, attracted more mosquitoes than the heavier Northern Cardinals, Cardinalis cardinalis (paired samples t-test, t=2.58, df=7, p=0.04). However, attractiveness did not differ substantially among bird species, and Gray Catbirds did not attract more mosquitoes than all other birds combined as a group. American Robins, Turdus migratorius (n=4) were comparable in attractiveness to other bird species, but not enough American Robins were captured for a comprehensive study of mosquito avian preference.