Fence Lizards Research Articles

Hypoxia Exposure Fine-Tunes Mitochondrial Function in Primary Dermal Fibroblasts Derived from Loggerhead Sea Turtles

Sea turtles are constantly exposed to changes in oxygen tension derived from extended breath-hold diving, but the cellular-level response to such changes is not completely understood. Most studies in hypoxia-tolerant reptiles have focused on the freshwater turtle Trachemys scripta, which responds to anoxia by inducing metabolic rewiring and preserving mitochondrial integrity, facilitating effcient mitochondrial respiration during reoxygenation. Here, we derived primary dermal fibroblasts from loggerhead sea turtles and western fence lizards to study the bioenergetic cellular response to hypoxia exposure in diving and non-diving reptiles. Cells from both species stained positive for the fibroblast markers vimentin and PDGFRb and responded to hypoxia exposure (0.1% O2) by accumulating HIF-1α. We measured oxygen consumption rates (OCR) in intact cells using Seahorse technology. We found that basal, maximal, and spare respiratory capacity was lower in lizard than in loggerhead cells at 26oC under normoxic (room air) conditions (p < 0.0001). Exposure to short (1h) and long-term hypoxia (24h) altered mitochondrial function in both species. Maximal and spare respiratory capacity were lower in loggerhead than in lizard cells (p < 0.01) after 1h hypoxia followed by 1h reoxygenation. In contrast, basal and maximal respiratory capacity was higher in loggerhead compared to lizard cells after 24h hypoxia (p < 0.04). Nonetheless, the overall OCR after 24h hypoxia in both species was lower than that observed after 1h hypoxia. Overall, there was a significant reduction in cellular respiration in both species from short-term to long-term hypoxia exposure. In lizard cells, basal OCR decreased by a factor of three while maximal OCR decreased by a factor of 4.5. In loggerhead cells, both the basal and maximal OCR decreased by a factor of two. Our results demonstrate species-specific regulation of cellular respiration after hypoxia exposure. Remarkably, loggerhead cells show a steady downregulation of cellular respiration in response to hypoxia exposure and faster recovery after extended hypoxia (24h) than lizard cells, aligning with the remarkable hypoxic tolerance exhibited by sea turtles, which can endure up to 7 hours of breath-holding underwater. Understanding the cellular mechanisms that drive hypoxic tolerance in diving animals can provide insights for treating human conditions characterized by hypoxia signaling. Ford Foundation. This is the full abstract presented at the American Physiology Summit 2024 meeting and is only available in HTML format. There are no additional versions or additional content available for this abstract. Physiology was not involved in the peer review process.

Cutaneous Evaporative Water Loss in Lizards Changes Immediately with Temperature.

AbstractCutaneous evaporative water loss (CEWL) makes up a significant portion of total evaporative water loss in many terrestrial vertebrates. CEWL changes on evolutionary and acclimatory timescales in response to temperature and humidity. However, the lability of CEWL on acute timescales is unknown. To examine this, we increased or decreased body temperatures of western fence lizards (Sceloporus occidentalis) over a 15-min period while continuously recording CEWL with a handheld evaporimeter. CEWL increased in response to heating and decreased in response to cooling on the order of seconds. However, CEWL was different between heating and cooling groups at a common body temperature. We observed the same positive relationship between CEWL and body temperature, as well as the difference in CEWL between treatments, for deceased lizards that we opportunistically measured. However, deceased lizards had more extreme CEWL values for any given body temperature and treatment. Overall, our results suggest that both structural traits and active physiological processes likely influence the rates and plasticity of CEWL.

Comparing fear responses of two lizard species across habitats varying in human impact

Abstract Animals that are successful in urban habitats often have reduced antipredator responses toward people (sometimes called “fear” responses). However, few studies test whether sympatric species differ in their responses to humans, which may explain differing sensitivities to urbanization. Here, we quantified the behavioral and physiological responses to humans in two lizard species, side-blotched lizards (Uta stansburiana) and western fence lizards (Sceloporus occidentalis), across three different habitat types that vary in human impact: natural habitats with low levels of human activity, natural habitats with high levels of human activity, and urban habitats. We found that side-blotched lizards had longer flight initiation distances, were found closer to a refuge, and were more likely to hide than fence lizards, behaviors that could indicate greater fearfulness. Both lizard species were found closer to a refuge and were also more likely to hide in the urban habitat than in the natural habitat with low human impact, which could represent adaptive behaviors for increased risks in urban areas (e.g. cats). Western fence lizards exhibited lower body sizes and conditions in the habitats with moderate and high levels of human activity, whereas these traits did not differ among habitats in side-blotched lizards. Baseline and stress-induced corticosterone concentrations did not differ across habitats for both species, suggesting that human-impacted habitats were not stressful or that lizards had undergone habituation-like processes in these habitats. Taken together, our results highlight the importance of standardized measurements across multiple species in the same habitats to understand differential responses to human-induced environmental change.

Tick-tock, racing the clock: parasitism is associated with decreased sprint performance in the Eastern fence lizard

Abstract Host–parasite relationships are important components of ecological systems that influence the evolution of both hosts and parasites. High levels of ectoparasitic infections can disrupt host homeostasis, causing adverse effects on health and performance. However, the effects of natural ectoparasite levels on host physiology are less understood, with most research designs implementing experimental or hormonal manipulations of hosts. In this study, we examined the interplay between tick parasitism and host characteristics on body condition and locomotor performance in Eastern fence lizards in natural settings. We found a higher frequency of tick infections in male lizards relative to females, with larger males being more likely to experience tick infection. Infected lizards had reduced locomotor performance, max sprint speed and 2-m run speed. Together this suggests an energetic trade-off between increased immune function and reduced locomotor performance, which is consistent with the immunocompetence–handicap hypothesis. Higher numbers of ticks on adult male lizards may be explained by age as well as the immunosuppressive effects of testosterone. Tick infection did not appear to reduce overall body condition of lizard hosts. Our findings shed light on the interplay between ectoparasitic infection, host characteristics, and locomotor performance under natural conditions. Such insights are crucial for understanding host–parasite dynamics and determining the trade-offs for hosts within ecological contexts.

Inter- and intrapopulation resource use variation of marine subsidized western fence lizards

Marine resource subsidies alter consumer dynamics of recipient populations in coastal systems. The response to these subsidies by generalist consumers is often not uniform, creating inter- and intrapopulation diet variation and niche diversification that may be intensified across heterogeneous landscapes. We sampled western fence lizards, Sceloporus occidentalis, from Puget Sound beaches and coastal and inland forest habitats, in addition to the lizards’ marine and terrestrial prey items to quantify marine and terrestrial resource use with stable isotope analysis and mixing models. Beach lizards had higher average δ13C and δ15N values compared to coastal and inland forest lizards, exhibiting a strong mixing line between marine and terrestrial prey items. Across five beach sites, lizard populations received 20–51% of their diet from marine resources, on average, with individual lizards ranging between 7 and 86% marine diet. The hillslope of the transition zone between marine and terrestrial environments at beach sites was positively associated with marine-based diets, as the steepest sloped beach sites had the highest percent marine diets. Within-beach variation in transition zone slope was positively correlated with the isotopic niche space of beach lizard populations. These results demonstrate that physiography of transitional landscapes can mediate resource flow between environments, and variable habitat topography promotes niche diversification within lizard populations. Marine resource subsidization of Puget Sound beach S. occidentalis populations may facilitate occupation of the northwesternmost edge of the species range. Shoreline restoration and driftwood beach habitat conservation are important to support the unique ecology of Puget Sound S. occidentalis.

The effects of early-life and intergenerational stress on the brain.

Stress experienced during ontogeny can have profound effects on the adult phenotype. However, stress can also be experienced intergenerationally, where an offspring's phenotype can be moulded by stress experienced by the parents. Although early-life and intergenerational stress can alter anatomy, physiology, and behaviour, nothing is known about how these stress contexts interact to affect the neural phenotype. Here, we examined how early-life and intergenerational stress affect the brain in eastern fence lizards (Sceloporus undulatus). Some lizard populations co-occur with predatory fire ants, and stress from fire ant attacks exerts intergenerational physiological and behavioural changes in lizards. However, it is unclear if intergenerational stress, or the interaction between intergenerational and early-life stress, modulates the brain. To test this, we captured gravid females from fire ant invaded and uninvaded populations, and subjected offspring to three early-life stress treatments: (1) fire ant attack, (2) corticosterone, or (3) a control. Corticosterone and fire ant attack decreased some aspects of the neural phenotype while population of origin and the interaction of early-life stress and population had no effects on the brain. These results suggest that early-life stressors may better predict adult brain variation than intergenerational stress in this species.

Effects of testosterone on urogenital tract morphology and androgen receptor expression in immature Eastern Fence lizards (Sceloporus undulatus)

In non-avian reptiles, the onset of sexual dimorphism of the major structures of the urogenital tract varies temporally relative to gonadal differentiation, more so than in other amniote lineages. In the current study, we used tonic-release implants to investigate the effects of exogenous testosterone (T) on postnatal development of the urogenital tract in juvenile Eastern Fence Lizards (Sceloporus undulatus) to better understand the mechanisms underlying the ontogeny of sexual differentiation in reptiles. We examined gonads, mesonephric kidneys and ducts (male reproductive tract primordia), paramesonephric ducts (oviduct primordia), sexual segments of the kidneys (SSKs), and hemiphalluses to determine which structures were sexually dimorphic independent of T treatment and which structures exhibited sexually dimorphic responses to T. To better understand tissue-level responsiveness to T treatment, we also characterized androgen receptor (AR) expression by immunohistochemistry. At approximately 4 months after hatching in control animals, gonads were well differentiated but quiescent; paramesonephric ducts had fully degenerated in males; mesonephric kidneys, mesonephric ducts, and SSKs remained sexually undifferentiated; and hemiphalluses could not be everted in either sex. Exogenous T caused enlargement, regionalization, and secretory activity of the mesonephric ducts and SSKs in both sexes; enlargement and regionalization of the oviducts in females; and enlargement of male hemipenes. The most responsive tissues exhibited moderate but diffuse staining for AR in control lizards and intense nuclear staining in T-treated lizards, suggestive of autoregulation of AR. The similarity between sexes in the responsiveness of the mesonephric ducts and SSK to T indicates an absence of sexually dimorphic organizational effects in these structures prior to treatment, which was initiated approximately 2 months after hatching. In contrast, the sex-specific responses in oviducts and hemipenes indicate that significant organization and/or differentiation had taken place prior to treatment.

Hydration and evaporative water loss of lizards change in response to temperature and humidity acclimation.

Testing acclimation plasticity informs our understanding of organismal physiology and applies to conservation management amidst our rapidly changing climate. While there is a wealth of research on the plasticity of thermal and hydric physiology in response to temperature acclimation, there is a comparative gap for research on acclimation to different hydric regimes, as well as the interaction between water and temperature. We sought to fill this gap by acclimating Western Fence Lizards (Sceloporus occidentalis) to experimental climate conditions (crossed design of Hot or Cool, Dry or Humid) for eight days, and measuring cutaneous evaporative water loss (CEWL), plasma osmolality, hematocrit, and body mass before and after acclimation. CEWL changed plastically in response to the different climates, with lizards acclimated to Hot Humid conditions experiencing the greatest increase in CEWL. Change in CEWL among individuals was negatively related to treatment vapor pressure deficit and positively related to treatment water vapor pressure. Plasma osmolality, hematocrit, and body mass all showed greater changes in response to temperature than to humidity or vapor pressure deficit. CEWL and plasma osmolality were positively related across treatment groups before acclimation and within treatment groups after acclimation, but the two variables showed different responses to acclimation, suggesting that they are interrelated but governed by different mechanisms. This study is among few that assess more than one metric of hydric physiology and that test the interactive effects of temperature and humidity. Such measurements will be essential for predictive models of activity and survival for animals under climate change.

Chigger mite (Eutrombicula alfreddugesi) ectoparasitism does not contribute to sex differences in growth rate in eastern fence lizards (Sceloporus undulatus).

Parasitism is nearly ubiquitous in animals and is frequently associated with fitness costs in host organisms, including reduced growth, foraging, and reproduction. In many species, males tend to be more heavily parasitized than females and thus may bear greater costs of parasitism. Sceloporus undulatus is a female-larger, sexually size dimorphic lizard species that is heavily parasitized by chigger mites (Eutrombicula alfreddugesi). In particular, the intensity of mite parasitism is higher in male than in female juveniles during the period of time when sex differences in growth rate lead to the development of sexual size dimorphism (SSD). Sex-biased differences in fitness costs of parasitism have been documented in other species. We investigated whether there are growth costs of mite ectoparasitism, at a time coinciding with sex differences in growth rate and the onset of SSD. If there are sex-biased growth costs of parasitism, then this could suggest a contribution to the development of SSD in S. undulatus. We measured growth and mite loads in two cohorts of unmanipulated, field-active yearlings by conducting descriptive mark-recapture studies during the activity seasons of 2016 and 2019. Yearling males had consistently higher mid-summer mite loads and consistently lower growth rates than females. However, we found that growth rate and body condition were independent of mite load in both sexes. Furthermore, growth rates and mite loads were higher in 2019 than in 2016. Our findings suggest that juveniles of S. undulatus are highly tolerant of chigger mites and that any costs imposed by mites may be at the expense of functions other than growth. We conclude that sex-biased mite ectoparasitism does not contribute to sex differences in growth rate and, therefore, does not contribute to the development of SSD.

Locomotor performance differs between foraging and predator escape but is not related to survival in hatchling lizards

Abstract Behaviour is influenced by many factors, including morphology, performance capabilities and the surrounding environment. Behavioural adjustments in response to these factors have important fitness consequences, and therefore behaviour should be considered when evaluating relationships among morphology, performance and fitness. We used hatchling fence lizards (Sceloporus undulatus) to test if sprinting speed and movement behaviours (e.g. latency to approach prey, number of pauses when fleeing or foraging) differ between foraging and anti-predator contexts and if these behaviours are associated with body size, locomotor performance capacity and survival. We quantified maximal sprint performance of individuals and evaluated their running speed in laboratory trials of foraging (moving towards crickets) and predator escape (response to a swooping avian predator). We show that hatchlings run at ~17–36% of total capacity when foraging. Although the most common anti-predator behaviour was to remain still, those that fled from the predator ran at full burst speed. However, foraging and escape speed/behaviours were not associated with body size or locomotor capacity. Hatchlings were subsequently released at our field site for a recapture study to assess phenotypic relationships with survival. We found no evidence that survival was influenced by performance or behaviour, but body size was positively related to survival in the field. These results highlight how seemingly important fitness-relevant behavioural and performance traits may not mediate the relationship between body size and survival.

Testing concordance and conflict in spatial replication of landscape genetics inferences.

The degree to which landscape genetics findings can be extrapolated to different areas of a species range is poorly understood. Here, we used a broadly distributed ectothermic lizard (Sceloporus occidentalis, Western Fence lizard) as a model species to evaluate the full role of topography, climate, vegetation, and roads on dispersal and genetic differentiation. We conducted landscape genetics analyses with a total of 119 individuals in five areas within the Sierra Nevada mountain range. Genetic distances calculated from thousands of ddRAD markers were used to optimize landscape resistance surfaces and infer the effects of landscape and topographic features on genetic connectivity. Across study areas, we found a great deal of consistency in the primary environmental gradients impacting genetic connectivity, along with some site-specific differences, and a range in the proportion of genetic variance explained by environmental factors across study sites. High-elevation colder areas were consistently found to be barriers to gene flow, as were areas of high ruggedness and slope. High temperature seasonality and high precipitation during the winter wet season also presented a substantial barrier to gene flow in a majority of study areas. The effect of other landscape variables on genetic differentiation was more idiosyncratic and depended on specific attributes at each site. Across study areas, canyon valleys were always implicated as facilitators to dispersal and key features linking populations and maintaining genetic connectivity, though the relative importance varied in different areas. We emphasize that spatial data layers are complex and multidimensional, and careful consideration of spatial data correlation structure and robust analytic frameworks will be critical to our continued understanding of spatial genetics processes.

Assembly of the largest squamate reference genome to date: The western fence lizard, Sceloporus occidentalis.

Spiny lizards (genus Sceloporus) have long served as important systems for studies of behavior, thermal physiology, dietary ecology, vector biology, speciation, and biogeography. The western fence lizard, Sceloporus occidentalis, is found across most of the major biogeographical regions in the western United States and northern Baja California, Mexico, inhabiting a wide range of habitats, from grassland to chaparral to open woodlands. As small ectotherms, Sceloporus lizards are particularly vulnerable to climate change, and S. occidentalis has also become an important system for studying the impacts of land use change and urbanization on small vertebrates. Here, we report a new reference genome assembly for S. occidentalis, as part of the California Conservation Genomics Project (CCGP). Consistent with the reference genomics strategy of the CCGP, we used Pacific Biosciences HiFi long reads and Hi-C chromatin-proximity sequencing technology to produce a de novo assembled genome. The assembly comprises a total of 608 scaffolds spanning 2,856 Mb, has a contig N50 of 18.9 Mb, a scaffold N50 of 98.4 Mb, and BUSCO completeness score of 98.1% based on the tetrapod gene set. This reference genome will be valuable for understanding ecological and evolutionary dynamics in S. occidentalis, the species status of the California endemic island fence lizard (S. becki), and the spectacular radiation of Sceloporus lizards.

Contextualized Display Behavior during Natural Interactions by Common Side-Blotched Lizards (Uta stansburiana, Squamata, Phrynosomatidae)

Many previous studies on competition focused on outcomes of competition and less on behavioral changes during or immediately following intra- or interspecific interactions. I performed 300 h of observation of three lizard species (Side-blotched Lizards, Uta stansburiana; Sagebrush Lizards, Sceloporus graciosus; Western Fence Lizards, Sceloporus occidentalis) and recorded 1,045 natural encounters, 403 of which were unconfounded by presence of another individual lizard. Despite potential competition amongst these species, I found little evidence that Uta modify perch characteristics or engage in aggressive interactions toward either of the heterospecific lizards. However, Uta do alter interspecific display behavior depending on whether interactions are intrasexual or intersexual. Moreover, Uta move shorter distances following interactions with S. graciosus but greater distances after interactions with S. occidentalis. Uta appear to reserve aggressive or more-intense interactions for intersexual intraspecific encounters, most of which are initiated by males. Together these findings suggest that these three lizard species are not broadly engaged in competitive interactions and that habitat characteristics or microhabitat preferences enable co-occurrence.

Recovery of western black-legged tick and vertebrate populations after a destructive wildfire in an intensively-studied woodland in northern California.

Despite increasing severity and frequency of wildfires, knowledge about how fire impacts the ecology of tick-borne pathogens is limited. In 2018, the River Fire burned a forest in the far-western U.S.A. where the ecology of tick-borne pathogens had been studied for decades. Forest structure, avifauna, large and small mammals, lizards, ticks, and tick-borne pathogens (Anaplasma phagocytophilum, Borrelia burgdorferi, Borrelia miyamotoi) were assessed after the wildfire in 2019 and 2020. Burning reduced canopy cover and eliminated the layer of thick leaf litter that hosted free-living ticks, which over time was replaced by forbs and grasses. Tick abundance and the vertebrate host community changed dramatically. Avian species adapted to cavity nesting became most prevalent, while the number of foliage-foraging species increased by 83% as vegetation regenerated. Nine mammalian species were observed on camera traps, including sentinel (black-tailed jackrabbits) and reservoir hosts (western gray squirrels) of B. burgdorferi. One Peromyscus sp. mouse was captured in 2019 but by 2020, numbers were rebounding (n=37), although tick infestations on rodents remained sparse (0.2/rodent). However, western fence lizards (n=19) hosted 8.6 ticks on average in 2020. Assays for pathogens found no B. miyamotoi in either questing or host-feeding ticks, A. phagocytophilum DNA in 4% (1/23) in 2019, and 17% (29/173) in 2020 for questing and host-feeding ticks combined, and B. burgdorferi DNA in just 1% of all ticks collected in 2020 (2/173). We conclude that a moderately severe wildfire can have dramatic impacts on the ecology of tick-borne pathogens, with changes posited to continue for multiple years.

Drought correlates with reduced infection complexity and possibly prevalence in a decades-long study of the lizard malaria parasite Plasmodium mexicanum.

Microparasites often exist as a collection of genetic 'clones' within a single host (termed multi-clonal, or complex, infections). Malaria parasites are no exception, with complex infections playing key roles in parasite ecology. Even so, we know little about what factors govern the distribution and abundance of complex infections in natural settings. Utilizing a natural dataset that spans more than 20 years, we examined the effects of drought conditions on infection complexity and prevalence in the lizard malaria parasite Plasmodium mexicanum and its vertebrate host, the western fence lizard, Sceloporus occidentalis. We analyzed data for 14,011 lizards sampled from ten sites over 34 years with an average infection rate of 16.2%. Infection complexity was assessed for 546 infected lizards sampled during the most recent 20 years. Our data illustrate significant, negative effects of drought-like conditions on infection complexity, with infection complexity expected to increase by a factor of 2.27 from the lowest to highest rainfall years. The relationship between rainfall and parasite prevalence is somewhat more ambiguous; when prevalence is modeled over the full range in years, a 50% increase in prevalence is predicted between the lowest and highest rainfall years, but this trend is not apparent or is reversed when data are analyzed over a shorter timeframe. To our knowledge, this is the first reported evidence for drought affecting the abundance of multi-clonal infections in malaria parasites. It is not yet clear what mechanism might connect drought with infection complexity, but the correlation we observed suggests that additional research on how drought influences parasite features like infection complexity, transmission rates and within-host competition may be worthwhile.

Sex and Season Affect Cortical Volumes in Free-Living Western Fence Lizards, Sceloporus occidentalis

The hippocampus plays an important role in spatial navigation and spatial learning across a variety of vertebrate species. Sex and seasonal differences in space use and behavior are known to affect hippocampal volume. Similarly, territoriality and differences in home range size are known to affect the volume of the reptile hippocampal homologues, the medial and dorsal cortices (MC, DC). However, studies have almost exclusively investigated males and little is known about sex or seasonal differences in MC and/or DC volumes in lizards. Here, we are the first to simultaneously examine sex and seasonal differences in MC and DC volumes in a wild lizard population. In Sceloporus occidentalis, males display territorial behaviors that are more pronounced during the breeding season. Given this sex difference in behavioral ecology, we expected males to have larger MC and/or DC volumes than females and for this difference to be most pronounced during the breeding season when territorial behavior is increased. Male and female S. occidentalis were captured from the wild during the breeding season and the post-breeding season and were sacrificed within 2 days of capture. Brains were collected and processed for histology. Cresyl-violet-stained sections were used to quantify brain region volumes. In these lizards, breeding females had larger DC volumes than breeding males and nonbreeding females. There was no sex or seasonal difference in MC volumes. Differences in spatial navigation in these lizards may involve aspects of spatial memory related to breeding other than territoriality that affect plasticity of the DC. This study highlights the importance of investigating sex differences and including females in studies of spatial ecology and neuroplasticity.

Biogeographic inferences across spatial and evolutionary scales.

The field of biogeography unites landscape genetics and phylogeography under a common conceptual framework. Landscape genetics traditionally focuses on recent-time, population-based, spatial genetics processes at small geographical scales, while phylogeography typically investigates deep past, lineage- and species-based processes at large geographical scales. Here, we evaluate the link between landscape genetics and phylogeographical methods using the western fence lizard (Sceloporus occidentalis) as a model species. First, we conducted replicated landscape genetics studies across several geographical scales to investigate how population genetics inferences change depending on the spatial extent of the study area. Then, we carried out a phylogeographical study of population structure at two evolutionary scales informed by inferences derived from landscape genetics results to identify concordance and conflict between these sets of methods. We found significant concordance in landscape genetics processes at all but the largest geographical scale. Phylogeographical results indicate major clades are restricted to distinct river drainages or distinct hydrological regions. At a more recent timescale, we find minor clades are restricted to single river canyons in the majority of cases, while the remainder of river canyons include samples from at most two clades. Overall, the broad-scale pattern implicating stream and river valleys as key features linking populations in the landscape genetics results, and high degree of clade specificity within major topographic subdivisions in the phylogeographical results, is consistent. As landscape genetics and phylogeography share many of the same objectives, synthesizing theory, models and methods between these fields will help bring about a better understanding of ecological and evolutionary processes structuring genetic variation across space and time.

Ecology and Epidemiology of Lyme Disease in Western North America

Lyme disease is the most common vector-borne disease in the United States and Canada. The causative agent of Lyme disease in North America is the spirochete Borrelia burgdorferi. In western North America, the primary vector of Borrelia burgdorferi is the western black-legged tick, Ixodes pacificus. Surveillance and modelling efforts indicate that I. pacificus is primarily found in coastal California, Oregon, Washington and the southern coastal regions of British Columbia However, infection rates with B. burgdorferi among I. pacificus ticks remain low, ranging from 0.6% to 9.9%. Lyme disease case numbers in western North America are also relatively low compared to eastern North America. Enzootic maintenance of B. burgdorferi by hosts in natural environments and climatic factors may influence Lyme disease risk. The borreliacidal western fence lizard, Sceloporus occidentalis, may contribute to the low infection rates observed in I. pacificus ticks, while the migratory nature of avian hosts can allow for long-distance tick dispersal. Moderately warm and moist environments and protection from sunlight define the suitable habitats of I. pacificus ticks. In this review, we discuss the ecology and epidemiology of Lyme disease in relation to I. pacificus, as well as the need for more studies in western North America.

Dilution Effect of the Refractory Lizard Host on the Transmission of the Lyme Disease Pathogen by Its Tick Vector in Northern California

A current model for Ixodes scapularis was revised and extended to represent the temperature dependent life cycle and host structure for Ixodes pacificus, the vector of the Lyme disease pathogen Borrelia burgdorferi in the Western United States. Parameters were adjusted to reflect data collected in Mendocino, CA. The hosts for I. pacificus include the Western Fence Lizard, which is not only an incompetent host for B. burgdorferi, but some studies claim that it eliminates it in ticks that feed on this lizard. The lizard host has been proposed as a reason for lower prevalence of Lyme disease in the West. This hypothesis was tested quantitatively by varying the number of lizard hosts. Model results suggest that the presence of refractory hosts produces a dilution effect for B. burgdorferi infection and is likely to be the reason for observed lower prevalence of Lyme disease in the Western United States.

13. A case of nocturnal activity in the diurnal southwestern fence lizard Scelopurus cowlesi

13. A case of nocturnal activity in the diurnal southwestern fence lizard Scelopurus cowlesi