Ecology of Ixodes pacificus Ticks and Associated Pathogens in the Western United States

Lyme disease is the most important vector-borne disease in the United States and is increasing in incidence and geographic range. In the Pacific west, the western black-legged tick, Ixodes pacificus Cooley and Kohls, 1943 is an important vector of the causative agent of Lyme disease, the spirochete, Borrelia burgdorferi. Ixodes pacificus life cycle is expected to be more than a year long, and all three stages (larva, nymph, and adult) overlap in spring. The optimal habitat consists of forest cover, cooler temperatures, and annual precipitation in the range of 200–500 mm. Therefore, the coastal areas of California, Oregon, and Washington are well suited for these ticks. Immature stages commonly parasitize Western fence lizards (Sceloporus occidentalis) and gray squirrels (Sciurus griseus), while adults often feed on deer mice (Peromyscus maniculatus) and black-tailed deer (Odocoileus h. columbianus). Ixodes pacificus carry several pathogens of human significance, such as Borrelia burgdorferi, Bartonella, and Rickettsiales. These pathogens are maintained in the environment by many hosts, including small mammals, birds, livestock, and domestic animals. Although a great deal of work has been carried out on Ixodes ticks and the pathogens they transmit, understanding I. pacificus ecology outside California still lags. Additionally, the dynamic vector–host–pathogen system means that new factors will continue to arise and shift the epidemiological patterns within specific areas. Here, we review the ecology of I. pacificus and the pathogens this tick is known to carry to identify gaps in our knowledge.

A lizard with two tales: what diversification within Sceloporus occidentalis teaches us about species formation

In 1859, Charles Darwin proposed that species are not fundamentally different from subspecies or the varieties from which they evolve. A century later, Dobzhansky (1958) suggested that many such lineages are ephemeral and are likely to revert differentiation through introgression (Fig. 1A); only a few evolve complete reproductive isolation and persist in sympatry. In this issue of Molecular Ecology, Bouzid et al. (2021) show how new analytical methods, when applied to genome data, allow us to more precisely determine whether or not species formation follows the paths outlined by Darwin and Dobzhansky (Fig. 1B). The authors study the diversification of the lizard Sceloporus occidentalis, finding a continuum of genetic interactions between the preservation of genetic identity to genetic merger, analogous to what is exemplified by ring species. In doing so, they teach us two tales on species formation: that lineages are fractal byproducts of evolutionary processes such as genetic drift and selection, and that lineages are often ephemeral and do not always progress into species. Studying ephemeral lineages like those in S. occidentalis allows us to capture divergence at its earliest stages, and potentially to determine the factors that allow lineages to remain distinct despite pervasive gene flow. These lineages thus serve as a natural laboratory to address long standing hypotheses on species formation.

First verified sighting of a Western Fence Lizard (Sceloporus occidentalis) in British Columbia, Canada

Western Fence Lizard (Sceloporus occidentalis) is known from Baja California, Mexico, north to north-central Washington State, including Puget Sound, where scattered populations extend from the Cherry Point area south to Tacoma and along the west side of Puget Sound to Port Townsend. On 6 June 2020, a single juvenile S. occidentalis was photographed in a Cloverdale area garden, Surrey, British Columbia, representing the first verified sighting of this species in Canada. No other S. occidentalis were sighted in the area, and we could not determine how the specimen entered the province.

The Lizard Gut Microbiome Changes with Temperature and Is Associated with Heat Tolerance

ABSTRACT Vertebrates harbor trillions of microorganisms in the gut, collectively termed the gut microbiota, which affect a wide range of host functions. Recent experiments in lab-reared vertebrates have shown that changes in environmental temperature can induce shifts in the gut microbiota, and in some cases these shifts have been shown to affect host thermal physiology. However, there is a lack of information about the effects of temperature on the gut microbiota of wild-caught vertebrates. Moreover, in ectotherms, which are particularly vulnerable to changing temperature regimens, the extent to which microbiota composition is shaped by temperature and associated with host thermal tolerance has not been investigated. To address these issues, we monitored the gut microbiota composition of wild-caught western fence lizards (Sceloporus occidentalis) experimentally exposed to a cool-to-warm temperature transition. Comparing experimentally exposed and control lizards indicated that warm temperatures altered and destabilized the composition of the S. occidentalis gut microbiota. Warming drove a significant reduction in the relative abundances of a clade of Firmicutes, a significant increase in the rate of compositional turnover in the gut microbiota within individual lizards, and increases in the abundances of bacteria from predicted pathogenic clades. In addition, the composition of the microbiota was significantly associated with the thermal tolerance of lizards measured at the end of the experiment. These results suggest that temperature can alter the lizard gut microbiota, with potential implications for the physiological performance and fitness of natural populations. IMPORTANCE Gut microbial communities affect their animal hosts in numerous ways, motivating investigations of the factors that shape the gut microbiota and the consequences of gut microbiota variation for host traits. In this study, we tested the effects of increases in environmental temperatures on the gut microbiota of fence lizards, a vertebrate ectotherm threatened by warming climates. By monitoring lizards and their gut microbes during an experimental temperature treatment, we showed that the warming altered and destabilized the lizard gut microbiota. Moreover, measuring thermal performance of lizard hosts at the end of the experiment indicated that the composition of the gut microbiota was associated with host thermal tolerance. These results indicate that warming temperatures can alter the gut microbiota of vertebrate ectotherms and suggest relationships between variation in the gut microbiota and the thermal physiology of natural host populations.

Establishment of brown anoles (Anolis sagrei) across a southern California county and potential interactions with a native lizard species

The brown anole, Anolis sagrei, is a native species to the Caribbean; however, A. sagrei has invaded multiple parts of the USA, including Florida, Louisiana, Hawai’i and more recently California. The biological impacts of A. sagrei invading California are currently unknown. Evidence from the invasion in Taiwan shows that they spread quickly and when immediate action is not taken eradication stops being a viable option. In Orange County, California, five urban sites, each less than 100 ha, were surveyed for an average of 49.2 min. Approximately 200 A. sagrei were seen and verified across all survey sites. The paucity of native lizards encountered during the surveys within these sites suggests little to no overlap between the dominant diurnal western fence lizard, Sceloporus occidentalis, and A. sagrei. This notable lack of overlap could indicate a potentially disturbing reality that A. sagrei are driving local extirpations of S. occidentalis.

Urban invaders are not bold risk-takers: a study of 3 invasive lizards in Southern California

Biological invasions threaten biodiversity worldwide, and therefore, understanding the traits of successful invaders could mitigate their spread. Many commonly invasive species do well in disturbed habitats, such as urban environments, and their abilities to effectively respond to disturbances could contribute to their invasiveness. Yet, there are noninvasive species that also do well in disturbed habitats. The question remains whether urban invaders behave differently in urban environments than noninvaders, which could suggest an “urban-exploiting” phenotype. In Southern California, the co-occurrence of invasive Italian wall lizards Podarcis siculus, brown anoles Anolis sagrei, and green anoles A. carolinensis, and native western fence lizards Sceloporus occidentalis offers an opportunity to test whether invasives exhibit consistent differences in risk-taking within human-altered habitats compared with a native species. We predicted that invasive lizards would exhibit more bold behavior by having shorter flight-initiation distances (FIDs) and by being found farther from a refuge (behaviors that would presumably maximize foraging in low-risk environments). Invasive populations had similar or longer FIDs, but were consistently found at distances closer to a refuge. Collectively, invasive lizards in urban habitats were not bolder than a native species. Reliance on nearby refuges might help species successfully invade urban habitats, and if a general pattern, may pose an added challenge in detecting or eliminating them.

Ecological variability is associated with functional trait diversity in the western fence lizard (Sceloporus occidentalis)

A major goal of evolutionary ecology is to understand the ways in which ecological variability has structured morphological diversity. The aim of this study was to examine intraspecific phenotypic variation in the western fence lizard (Sceloporus occidentalis) for functional traits previously shown to be linked to variation across latitude and climate at a genus-wide level. We found that body size in S. occidentalis was negatively related to minimum temperatures during the coldest month, potentially implicating overwintering survival as a mechanism leading to this relationship. We also demonstrated that scale size in S. occidentalis was negatively associated with aridity, which is consistent with several other studies in squamate reptiles and the known functional role of scale size in moderating evaporative water loss. However, contrary to predictions of the thermal melanism hypothesis, we found no association between lizard colour and temperature during the active season. Overall, our results are largely at odds with interspecific patterns examining the relationship between these traits and environmental conditions. It is unclear to what degree intraspecific trait variation generally agrees with inferences made at the interspecific level. In any case, more studies at the intraspecific level are needed to resolve this question.