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Abstract BackgroundAs habitat fragmentation increases, ecological processes, including patterns of vector-borne pathogen prevalence, will likely be disrupted, but ongoing investigations are necessary to examine this relationship. Here, we report the differences in the prevalence of Lyme disease (Borrelia burgdorferisensu lato, s.l.) and haemoproteosis (Haemoproteusspp.) pathogens in avian populations of a fragmented habitat.B. burgdorferis.l. is a generalist pathogen that is transmitted byIxodes pacificusvectors in California, andHaemoproteusis an avian parasite transmitted byCulicoidesvectors. MethodsTo determine whether biotic (avian and mammalian abundance) or abiotic characteristics (patch size and water availability) correlated with infection prevalence change, we screened 176 birds sampled across seven sites in oak woodland habitat in northern California. ResultsWhile biotic factors correlated with an increase in both pathogens, infection prevalence ofHaemoproteusspp. was only associated with individual-level traits, specifically foraging substrate and diet, andB. burgdorferis.l. was associated with community-level characteristics, both total mammal and, specifically, rodent abundance. Proximity to water was the only abiotic factor found to be significant for both pathogens and reinforces the importance of water availability for transmission cycles. Larger patch sizes did not significantly affect infection prevalence ofHaemoproteus,but did increase the prevalence ofB. burgdorferi. ConclusionsThese results highlight that while environmental factors (specifically habitat fragmentation) have a limited role in vector-borne pathogen prevalence, the indirect impact to biotic factors (community composition) can have consequences for bothHaemoproteusandB. burgdorferiprevalence in birds. Given the pervasiveness of habitat fragmentation, our results are of broad significance. Graphical abstract 

Abstract Pacific Coast tick fever is a recently described zoonotic disease in California caused by a spotted fever group rickettsia, Rickettsia rickettsii subsp. californica (formerly Rickettsia 364D) and transmitted by the Pacific Coast tick, Dermacentor occidentalis. Like many emerging vector-borne diseases, knowledge regarding the transmission cycle, contribution from potential amplifying hosts, and geographic distribution of R. rickettsii californica is limited. We paired molecular analysis with comparative spatial niche modeling to identify vertebrate hosts potentially involved in the transmission cycle of this pathogen. We identified R. rickettsii californica DNA in three mammal species (Otospermophilus beecheyi, Lepus californicus, and Sylvilagus audubonii). This is the first record of R. rickettsii californica detected in mammals and may indicate potential amplifying hosts for this human pathogen. Species niche modeling of uninfected and infected D. occidentalis identified areas of high suitability along the coast and Sierra Nevada foothills of California. These findings support the hypothesis that amplifying host(s) may support higher infection prevalence in the infected tick regions compared to other parts of the tick’s range. Potential host species distribution models (SDMs) were constructed from museum records and niche overlap statistics were used to compare habitat suitability with R. rickettsii californica-infected tick SDMs. We found higher than null overlap of infected ticks with California ground squirrels (O. beecheyii) and trending, but nonsignificant, overlap with two lagomorph species. Pairing molecular and niche modeling may be a useful approach to identify species that are involved in the maintenance of emerging tick-borne zoonoses. 

Abstract Changing climate has driven shifts in species phenology, influencing a range of ecological interactions from plant–pollinator to consumer–resource. Phenological changes in host–parasite systems have implications for pathogen transmission dynamics. The seasonal timing, or phenology, of peak larval and nymphal tick abundance is an important driver of tick‐borne pathogen prevalence through its effect on cohort‐to‐cohort transmission. Tick phenology is tightly linked to climatic factors such as temperature and humidity. Thus, variation in climate within and across regions could lead to differences in phenological patterns. These differences may explain regional variation in tick‐borne pathogen prevalence of the Lyme disease‐causingBorreliabacteria in vector populations in the United States. For example, one factor thought to contribute to high Lyme disease prevalence in ticks in the eastern United States is the asynchronous phenology of ticks there, where potentially infected nymphal ticks emerge earlier in the season than uninfected larval ticks. This allows the infected nymphal ticks to transmit the pathogen to hosts that are subsequently fed upon by the next generation of larval ticks. In contrast, in the western United States where Lyme disease prevalence is generally much lower, tick phenology is thought to be more synchronous with uninfected larvae emerging slightly before, or at the same time as, potentially infected nymphs, reducing horizontal transmission potential. Sampling larval and nymphal ticks, and their host‐feeding phenology, both across large spatial gradients and through time, is challenging, which hampers attempts to conduct detailed studies of phenology to link it with pathogen prevalence. In this study, we demonstrate through intensive within‐season sampling that the relative abundance and seasonality of larval and nymphal ticks are highly variable along a latitudinal gradient and likely reflect the variable climate in the far western United States with potential consequences for pathogen transmission. We find that feeding patterns were variable and synchronous feeding of juvenile ticks on key blood meal hosts was associated with mean temperature. By characterizing within‐season phenological patterns of the Lyme disease vector throughout a climatically heterogeneous region, we can begin to identify areas with high potential for tick‐borne disease risk and underlying mechanisms at a finer scale. 

Abstract Pathogens utilize different modes of transmission to maximize transmission success. In vector‐borne disease systems, both vertical and horizontal modes of transmission are common, but the relative contribution of these modes is not well understood but may be determined by host genetics, physiology, or environmental conditions. This study focuses on an emerging tick‐borne relapsing fever pathogen,Borrelia miyamotoi, that can be transmitted both vertically and horizontally. The enzootic cycle of this pathogen has not been described in the western USA where it was recently found in the tick species,Ixodes pacificus. Our field surveys found that all three life stages ofI. pacificuscarry the pathogen, and therefore, all stages pose some level of disease risk to humans. The prevalence of infection increases with each life stage suggesting that horizontal transmission is important in the persistence of this pathogen in the enzootic cycle. In support of this finding, we found that small mammal hosts that are frequently parasitized by juvenile stages ofI. pacificuswere infected withB. miyamotoiand may therefore function as a source of horizontal transmission and enzootic maintenance of this disease. Our data show that in the western USAB. miyamotoiis maintained in natural populations by both transovarial transmission and transmission from blood meal hosts and that synchronous phenology of juvenile stages ofI. pacificusmay facilitate the transmission dynamics ofB. miyamotoiand other vertically transmitted, vector‐borne pathogens. 

Habitat loss and forest fragmentation are often linked to increased pathogen transmission, but the extent to which habitat isolation and landscape connectivity affect disease dynamics through movement of disease vectors and reservoir hosts has not been well examined. Tick-borne diseases are the most prevalent vector-borne diseases in the United States and on the West Coast,Ixodes pacificusis one of the most epidemiologically important vectors. We investigated the impacts of habitat fragmentation on pathogens transmitted byI. pacificusand sought to disentangle the effects of wildlife communities and landscape metrics predictive of pathogen diversity, prevalence and distribution. We collected pathogen data for four co-occurring bacteria transmitted byI. pacificusand measured wildlife parameters. We also used spatial data and cost-distance analysis integrating expert opinions to assess landscape metrics of habitat fragmentation. We found that landscape metrics were significant predictors of tick density and pathogen prevalence. However, wildlife variables were essential when predicting the prevalence and distribution of pathogens reliant on wildlife reservoir hosts for maintenance. We found that landscape structure was an informative predictor of tick-borne pathogen richness in an urban matrix. Our work highlights the implications of large-scale land management on human disease risk. 

Wildlife species are often heavily parasitized by multiple infections simultaneously. Yet research on sylvatic transmission cycles, tend to focus on host interactions with a single parasite and neglects the influence of co- infections by other pathogens and parasites. Co-infections between macro-parasites and micro-parasites can alter mechanisms that regulate pathogenesis and are important for understanding disease emergence and dy- namics. Wildlife rodent hosts in the Lyme disease system are infected with macro-parasites (i.e., ticks and hel- minths) and micro-parasites (i.e., Borrelia spp.), however, there has not been a study that investigates the interaction of all three parasites (i.e., I. pacificus, Borrelia spp., and helminths) and how these co-infections impact prevalence of micro-parasites. We live-trapped rodents in ten sites in northern California to collect feces, blood, ear tissue, and attached ticks. These samples were used to test for infection status of Borrelia species (i.e., micro- parasite), and describe the burden of ticks and helminths (i.e., macro-parasites). We found that some rodent hosts were co-infected with all three parasites, however, the burden or presence of concurrent macro-parasites were not associated with Borrelia infections. For macro-parasites, we found that tick burdens were positively associ- ated with rodent Shannon diversity while negatively associated with predator diversity, whereas helminth burdens were not significantly associated with any host community metric. Ticks and tick-borne pathogens are associated with rodent host diversity, predator diversity, and abiotic factors. However, it is still unknown what factors helminths are associated with on the community level. Understanding the mechanisms that influence co- infections of multiple types of parasites within and across hosts is an increasingly critical component of characterizing zoonotic disease transmission and maintenance. 

Abstract The western black-legged tick (Ixodes pacificus) is the most frequently identified human-biting tick species in the western United States and the principal vector of at least three recognized bacterial pathogens of humans. A potentially pathogenic Rickettsia species, first described in 1978 and recently characterized as a novel transitional group agent designated as Rickettsia tillamookensis, also exists among populations of I. pacificus, although the distribution and frequency of this agent are poorly known. We evaluated DNA extracts from 348 host-seeking I. pacificus nymphs collected from 9 locations in five California counties, and from 916 I. pacificus adults collected from 24 locations in 13 counties, by using a real-time PCR designed specifically to detect DNA of R. tillamookensis. DNA of R. tillamookensis was detected in 10 (2.9%) nymphs (95% CI: 1.6–5.2%) and 17 (1.9%) adults (95% CI: 1.2–3.0%) from 11 counties of northern California. Although site-specific infection rates varied greatly, frequencies of infection remained consistently low when aggregated by stage, sex, habitat type, or geographical region. Four novel isolates of R. tillamookensis were cultivated in Vero E6 cells from individual adult ticks collected from Alameda, Nevada, and Yolo counties. Four historical isolates, serotyped previously as ‘Tillamook-like’ strains over 40 yr ago, were revived from long-term storage in liquid nitrogen and confirmed subsequently by molecular methods as isolates of R. tillamookensis. The potential public health impact of R. tillamookensis requires further investigation. 

A vector's susceptibility and ability to transmit a pathogen—termed vector competency—determines disease outcomes, yet the ecological factors influencing tick vector competency remain largely unknown. Ixodes pacificus, the tick vector of Borrelia burgdorferi (Bb) in the western U.S., feeds on rodents, birds, and lizards. Rodents and birds are reservoirs for Bb and infect juvenile ticks, while lizards are refractory to Bb and cannot infect feeding ticks. Additionally, the lizard bloodmeal contains borreliacidal properties, clearing previously infected feeding ticks of their Bb infection. Despite I. pacificus feeding on a range of hosts, it is undetermined how the host identity of the larval bloodmeal affects future nymphal vector competency. We experimentally evaluate the influence of larval host bloodmeal on Bb acquisition by nymphal I. pacificus. Larval I. pacificus were fed on either lizards or mice and after molting, nymphs were fed on Bb-infected mice. We found that lizard-fed larvae were significantly more likely to become infected with Bb during their next bloodmeal than mouse-fed larvae. We also conducted the first RNA-seq analysis on whole-bodied I. pacificus and found significant upregulation of tick antioxidants and antimicrobial peptides in the lizard-fed group. Our results indicate that the lizard bloodmeal significantly alters vector competency and gene regulation in ticks, highlighting the importance of host bloodmeal identity in vector-borne disease transmission and upends prior notions about the role of lizards in Lyme disease community ecology. 

Globally, zoonotic vector-borne diseases are on the rise and understanding their complex transmission cycles is pertinent to mitigating disease risk. In North America, Lyme disease is the most commonly reported vector-borne disease and is caused by transmission of Borrelia burgdorferi sensu lato (s.l.) from Ixodes spp. ticks to a diverse group of vertebrate hosts. Small mammal reservoir hosts are primarily responsible for maintenance of B. burgdorferi s.l. across the United States. Never- theless, birds can also be parasitized by ticks and are capable of infection with B. burgdorferi s.l. but their role in B. burgdorferi s.l. transmission dynamics is understudied. Birds could be important in both the maintenance and spread of B. burgdorferi s.l. and ticks because of their high mobility and shared habitat with important mammalian reservoir hosts. This study aims to better understand the role of avian hosts in tick-borne zoonotic disease transmission cycles in the western United States. We surveyed birds, mammals, and ticks at nine sites in northern California for B. burgdorferi s.l. infection and collected data on other metrics of host community composition such as abundance and diversity of birds, small mammals, lizards, predators, and ticks. We found 22.8% of birds infected with B. burgdorferi s.l. and that the likelihood of avian B. burgdorferi s.l. infection was significantly associated with local host community composition and pathogen prevalence in California. Addition- ally, we found an average tick burden of 0.22 ticks per bird across all species. Predator and lizard abundances were significant predictors of avian tick infestation. These results indicate that birds are relevant hosts in the local B. burgdorferi s.l. transmission cycle in the western United States and quantifying their role in the spread and maintenance of Lyme disease requires further research.