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Abstract BackgroundEctothermic arthropods, like ticks, are sensitive indicators of environmental changes, and their seasonality plays a critical role in the dynamics of tick-borne disease in a warming world. Juvenile tick phenology, which influences pathogen transmission, may vary across climates, with longer tick seasons in cooler climates potentially amplifying transmission. However, assessing juvenile tick phenology is challenging in arid climates because ticks spend less time seeking for blood meals (i.e. questing) due to desiccation pressures. As a result, traditional collection methods like dragging or flagging are less effective. To improve our understanding of juvenile tick seasonality across a latitudinal gradient, we examinedIxodes pacificusphenology on lizards, the primary juvenile tick host in California, and explored how climate factors influence phenological patterns. MethodsBetween 2013 and 2022, ticks were removed from 1527 lizards at 45 locations during peak tick season (March–June). Tick counts were categorized by life stage (larvae and nymphs) and linked with remotely sensed climate data, including monthly maximum temperature, specific humidity and Palmer Drought Severity Index (PDSI). Juvenile phenology metrics, including tick abundances on lizards, Julian date of peak mean abundance and temporal overlap between larval and nymphal populations, were analyzed along a latitudinal gradient. Generalized additive models (GAMs) were applied to assess climate-associated variation in juvenile abundance on lizards. ResultsMean tick abundance per lizard ranged from 0.17 to 47.21 across locations, with the highest abundance in the San Francisco Bay Area and lowest in Los Angeles, where more lizards had zero ticks attached. In the San Francisco Bay Area, peak nymphal abundance occurred 25 days earlier than peak larval abundance. Temporal overlap between larval and nymphal stages at a given location varied regionally, with northern areas showing higher overlap, possibly due to the bimodal seasonality of nymphs. We found that locations with higher temperatures and increased drought stress were linked to lower tick abundances, although the magnitude of these effects depended on regional location. ConclusionsOur study, which compiled 10 years of data, reveals significant regional variation in juvenileI. pacificusphenology across California, including differences in abundance, peak timing, and temporal overlap. These findings highlight the influence of local climate on tick seasonality, with implications for tick-borne disease dynamics in a changing climate. Graphical Abstract 

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 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. 

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. 

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. 

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 Large‐bodied wild ungulates are declining worldwide, while domestic livestock continue to increase in abundance. Such changes in large herbivore communities should have strong effects on the control of ticks and tick‐borne disease as they can indirectly modify habitat and directly serve as final hosts for ticks' lifecycles. Numerous studies have now linked changing ungulate communities to changes in tick populations and disease risk. However, the effects of changing large herbivore communities are variable across studies, and the effect of climate as a mediating factor of this variation remains poorly understood. Also, studies to date have largely focused on wildlife loss without considering the extent to which livestock additions may alter tick populations, even though livestock replacement of wildlife is the global norm. In this study, we used a large‐scale exclosure experiment replicated along a topo‐climatic gradient to examine the effects on tick populations of both large herbivore removal and livestock additions. We found that while questing ticks increased modestly, by 21%, when large herbivores were removed from a system they decreased more substantially, by 50%, when livestock (in the form of cattle) were added. Importantly, in addition to the direct effects of climate on tick populations, climate also mediates the effect of ungulates on questing tick density. Particularly, the addition of livestock under the most arid conditions decreased tick presence, likely due to changes in ground‐level microclimates away from those beneficial to ticks. Overall, the work contributes to our understanding of tick population responses to globally common human‐induced rangeland alterations under the concurrent effects of climate change. 

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.