Sceloporus Occidentalis Research Articles

Thermal tolerance and Hsp70 expression in the Western fence lizard,Sceloporus occidentalis: Geographic variation in measures of acute stress and their role in selection on sprint speeds.

The paradigm of local adaptation to temperature has been extensively studied in the laboratory as well as in the field. However, many studies have exclusively examined invertebrate species which are often restricted to thermally isolated microhabitats. In this study, we examined the potential for temperature adaptation in the western fence lizard, Sceloporus occidentalis, which occurs in the western United States across thermally heterogeneous habitats. We collected lizards from sites paired in altitude and latitude (four sites total) and measured thermal tolerance (CTMax and CTMin) as well as expression of Hsp70, a molecular chaperone produced in response to thermal stress. In addition, we measured sprint speed of lizards before and after exposure to an experimental heat stress (or control treatment). We found that CTMax increased at all sites during the course of the summer season and CTMin decreased during the same time period but only at some sites. We also found that all lizards exposed to a heat treatment suffered reduced sprint speeds relative to control animals but the magnitude of the decrease depended upon the original site of capture. Finally, across all sites, increased expression of Hsp70 was correlated with reduced sprint speeds. These results indicate the potential for temperature adaptation to act within these populations and may lead to locally adapted populations of S. occidentalis.

Reptile Infection with Anaplasma phagocytophilum, the Causative Agent of Granulocytic Anaplasmosis

Granulocytic anaplasmosis (GA) is a potentially fatal tick-borne rickettsial disease that occurs sporadically in the far western United States. We evaluated the prevalence of Anaplasma phagocytophilum in multiple species of lizards and snakes from enzootic sites in northern California, described the infestation prevalence of its tick vector Ixodes pacificus on reptiles, and conducted an experimental challenge of western fence lizards (Sceloporus occidentalis) and Pacific gopher snakes (Pituophis catenifer) with A. phagocytophilum delivered via needle inoculation or tick bite. Both serologically and polymerase-chain reaction (PCR)-positive lizards (seroprevalence = 10.8%, PCR prevalence = 10.2%) and snakes (seroprevalence = 5.8%, PCR prevalence = 11.7%) were detected among wild-caught animals. A DNA sequence of the A. phagocytophilum groESL gene from a PCR-positive snake was 100% homologous to that of the human-derived A. phagocytophilum. Experimental attempts to infect naïve animals were unsuccessful for snakes (n = 2), but 1 of 12 lizards became infected for 1 wk only by tick bite. Xenodiagnostic I. pacificus larvae that fed on a PCR-positive lizard did not acquire or transmit rickettsiae. Our findings suggest that lizards and snakes are exposed to A. phagocytophilum by infected ticks, but that they do not serve as primary reservoir hosts of this rickettsia.

Feeding Preferences of the Immature Stages of Three Western North American Ixodid Ticks (Acari) for Avian, Reptilian, or Rodent Hosts

Larval and nymphal Ixodes pacificus Cooley and Kohls, I. (Ixodes) jellisoni Cooley and Kohls, and Dermacentor occidentalis Marx were tested for host preference when simultaneously presented with a deer mouse (Peromyscus maniculatus Wagner), California kangaroo rat (Dipodomys californicus Merriam), western fence lizard (Sceloporus occidentalis Baird and Girard), and California towhee (Pipilo crissalis Vigors) in an experimental apparatus. Differences were observed in the preferences among the three species and between life stages. More larvae of all species approached and contacted hosts than did nymphs. Subadult I. pacificus entered all host-containing chambers in the highest numbers and remained on lizards most often after contact. Subadult I. jellisoni entered and remained in the chambers containing kangaroo rats, while rejecting mice, lizards, and birds as hosts. Subadult D. occidentalis most frequently entered rodent-containing chambers and contacted these hosts. After overnight exposure to all nonavian hosts, only I. pacificus parasitized and fed successfully on all three animals. I. jellisoni fed only on kangaroo rats and D. occidentalis fed only on rodents. Molting success ranged from approximately 66 to 95% among tick species and stages. We concluded that, under laboratory conditions, I. pacificus larvae and nymphs prefer western fence lizards, but also will parasitize rodents. Dermacentor occidentalis immatures use deer mice and kangaroo rats similarly, whereas I. jellisoni subadults exclusively parasitize kangaroo rats. California towhees are considerably less attractive as hosts for these three ticks. These host preferences are consistent with what is known about the natural feeding habits of all three ticks.

Caudal Movements in Western Fence Lizards (Sceloporus occidentalis) Prior to Attempted Prey Capture

Many snakes employ tail movements to attract prey, but this behavior is rare in lizards. Use of caudal movements to distract prey, however, is rare in all squamates. Recently, caudal movements in a Sceloporus occidentalis individual immediately prior to attempted prey capture were documented. We supplement this sole documentation of caudal movements in S. occidentalis with additional field observations. Additionally, video footage of the behavior was recorded on 2 occasions. From these additional observations, we hypothesize that this behavior might serve to distract prey.

Toxicity of oral exposure to 2,4,6‐trinitrotoluene in the western fence lizard (Sceloporus occidentalis)

Contamination of the soil with the explosive 2,4,6-trinitrotoluene (TNT) has been found at military sites, many of which are habitats used by reptiles. To provide data useful in assessing ecological risk for reptilian species, acute, subacute, and subchronic oral toxicity studies were conducted with the western fence lizard (Sceloporus occidentalis). Oral median lethal dose (LD50) values for TNT in corn oil were 1,038 and 1,579 mg/kg of body weight for male and female lizards, respectively. Overt signs of toxicity included chromaturia, abdominal enlargement, and tremors. A 14-d subacute study followed in which male lizards were orally dosed with TNT (corn oil) at 0, 33, 66, 132, 263, 525, and 1,050 mg/kg of body weight each day. Clinical signs of toxicity, while similar to the LD50 study, were more subtle and noted in lizards receiving TNT amounts of at least 66 mg/kg/d. Chromaturia was an early consistent sign, often preceding the onset of adverse effects. Male lizards in the 60-d subchronic study were dosed at 0, 3, 15, 25, 35, and 45 mg/kg/d with nearly complete survival (>90%) for lizards in all treatments. Changes in food consumption and body weight were observed at 35 and 45 mg/kg/d. Alterations in hematological end points; blood chemistries (albumin, total protein, alkaline phosphatase, calcium); kidney, spleen, and liver weights; and adverse histopathology were observed in lizards exposed at 25 to 45 mg/kg/d. Testosterone concentration, sperm count, and motility were variable between treatments. Although not significant, incidences of hypospermia and testicular atrophy were observed in some individuals. Together, these data suggest a lowest-observed-adverse effect level of 25 mg/kg/d and a no-observed-adverse effect level of 15 mg/kg/d in S. occidentalis.

Estimating maximum performance: effects of intraindividual variation

Researchers often estimate the performance capabilities of animals using a small number of trials per individual. This procedure inevitably underestimates maximum performance, but few studies have examined the magnitude of this effect. In this study we explored the effects of intraindividual variation and individual sample size on the estimation of locomotor performance parameters. We measured sprint speed of the lizard Sceloporus occidentalis at two temperatures (20 degrees C and 35 degrees C), obtaining 20 measurements per individual. Speed did not vary temporally, indicating no training or fatigue effects. About 50% of the overall variation in speed at each temperature was due to intraindividual variation. While speed was repeatable, repeatability decreased slightly with increasing separation between trials. Speeds at 20 degrees C and 35 degrees C were positively correlated, indicating repeatability across temperatures as well. We performed statistical sampling experiments in which we randomly drew a subset of each individual's full set of 20 trials. As expected, the sample's maximum speed increased with the number of trials per individual; for example, five trials yielded an estimate averaging 89% of the true maximum. The number of trials also influenced the sample correlation between mean speeds at 20 degrees C and 35 degrees C; for example, five trials yielded a correlation coefficient averaging 90% of the true correlation. Therefore, intraindividual variation caused underestimation of maximal speed and the correlation between speeds across temperatures. These biases declined as the number of trials per individual increased, and depended on the magnitude of intraindividual variation, as illustrated by running sampling experiments that used modified data sets.

The effects of prey species on food conversion efficiency and growth of an insectivorous lizard

Little is known about the effects of different prey species on lizard growth. We conducted a 6-week study to determine the relative effects of prey species on growth parameters of hatchling western fence lizards, Sceloporus occidentalis. Lizards were fed house cricket nymphs, Acheta domesticus, or mealworm larvae, Tenebrio molitor. The effects of prey species on growth were determined by measuring prey consumption, gross conversion efficiency of food [gain in mass (g)/food consumed (g)], gain in mass, and gain in snout-vent length. Lizards grew well on both the prey species. However, lizards that fed on crickets consumed a significantly higher percentage of their body mass per day than those fed mealworms. Nevertheless, lizards that consumed mealworms ingested significantly more metabolizable energy, had significantly higher food conversion efficiencies, significantly higher daily gains in mass, and significantly greater total growth in mass than lizards that fed on crickets.

Dose‐related effects following oral exposure of 2,4‐dinitrotoluene on the western fence lizard, Sceloporus occidentalis

2,4-dintitrotoluene (2,4-DNT) is an explosive frequently found in the soil of military installations. Because reptiles can be common on these sites, ecological risk assessments for compounds such as 2,4-DNT could be improved with toxicity data specific to reptiles. Western fence lizards, Sceloporus occidentalis, were used to develop a laboratory toxicity model for reptiles. A hierarchical approach was used; acute to subchronic studies were conducted to provide toxicity data relevant to short- and long-term exposures. First, a modified median lethal dose (LD50) study was conducted on male and female lizards using a stage-wise probit model. The LD50 was 577 mg/kg for female and 380 mg/kg for male lizards. Subsequently, a subacute experiment was conducted to further assess 2,4-DNT toxicity to male lizards and to define exposure levels for a longer term, subchronic study. The subchronic study was conducted for 60 consecutive days; male lizards were exposed to 0, 9, 15, 25, 42, 70 mg/kg/d. Dose-dependent mortality was observed in the three highest dose groups (25, 42, and 70 mg/kg/d); all other animals survived the study duration. Benchmark dose model calculations based on mortality indicated a 5% effect level of 15.8 mg/kg/d. At study termination, a gross necropsy was performed, organ weights were taken, and blood was collected for clinical and hematological analysis. Body weight, kidney weight, food consumption, postdose observations, and blood chemistries all were found to be significantly different from controls at doses above 9 mg/kg/d. Also, preliminary results suggest behavioral observations, and reduced food consumption may be a sensitive indicator of toxicity. The present study indicates Sceloporus occidentalis is suitable for evaluating toxicity of compounds to reptilian species.

Effect of exogenous corticosterone on respiration in a reptile

Release of glucocorticoids (GCs) enables organisms to meet energy requirements during stressful situations by regulating intermediary metabolism. In the absence of compensatory mechanisms, increased metabolic activity (e.g., protein catabolism, lipolysis, and gluconeogenesis) should translate to increases in whole animal metabolism, and therefore energy expenditures, by organisms. However, to our knowledge, no study has estimated the total energy cost of elevated plasma GCs in any organism. Here we evaluated the effect of exogenous corticosterone (CORT) on metabolism in captive western fence lizards ( Sceloporus occidentalis) by conducting two experiments. In experiment I we determined the dynamics of plasma CORT concentrations resulting from CORT injections. In experiment II we frequently measured changes in respiration for 24 h before and after CORT injection. Injection of 0.025 μg/g (low CORT) and 0.40 μg/g CORT (high CORT) produced up to 26-fold increases in plasma CORT in lizards 3 h following injection compared to baseline levels. Plasma CORT concentrations returned to baseline levels 6 h after injection. CORT increased metabolism in both low and high CORT lizards (up to 121% and 245%, respectively) during the first 7.5 h after CORT injections compared to pre-treatment trials. Respiration returned to baseline rates 7.5 h after CORT administration. A surprising finding was that although high CORT males achieved higher plasma CORT concentrations than high CORT females, the metabolic response of high CORT females was 30% greater than high CORT males. Our results suggest that GC-induced changes in respiration may be important for understanding the overall energetic implications of stress.

Reproductive and thyroid hormone profiles in captive Western fence lizards (Sceloporus occidentalis) after a period of brumation

Seasonal fluctuation in serum concentrations of sex steroid (testosterone [T] and 17beta-estradiol [E(2)]) and thyroid (triiodothyronine [T(3)] and thyroxine [T(4)]) hormones was determined in captive Western fence lizards (Sceloporus occidentalis). Samples were collected from male and female breeding pairs weekly for a 4-month period after their emergence from artificial brumation. Circulating levels of E(2) corresponded with the expected vitellogenic and ovulatory cycles in females, and surprisingly, E(2) in males followed a similar pattern, indicating a possible role in breeding behavior. Serum T was elevated in male lizards for the first 6 weeks after emergence from brumation, possibly related to an increase in the onset of active spermatogenesis. Thyroid hormones showed little cyclical activity throughout the breeding period, with the exception of small increases of T(3) at weeks 8 and 16, possibly implying an active role of this hormone with ovulation in females. Overall, these baseline hormone data are not only useful in developing this animal as a laboratory reptile model for assessment of endocrine-mediated toxicity, but also of value for understanding herpetological endocrinology and for application in the conservation of threatened species. Zoo Biol 27:36-48, 2008. (c) 2007 Wiley-Liss, Inc.

Fluctuating Asymmetry as an Indicator of Environmental Stress From Off‐Highway Vehicles

Abstract:With human activities increasingly impacting natural resources in relatively remote locations, there is a need for simple and efficient methods to explore the ecological consequences of these activities. Little is understood about the influences of off‐highway vehicle (OHV) use on wildlife populations. We examined the effect of OHV activity on developmental instability in a phrynosomatid lizard (i.e., western fence lizard [Sceloporus occidentalis]) in the western Great Basin, USA. We measured fluctuating asymmetry (FA) of bilateral head‐scale patterns in populations of lizards at 3 OHV and 3 non‐OHV sites. Fluctuating asymmetry was higher at OHV sites relative to non‐OHV sites, supporting the idea that OHV activity can stress wildlife populations. We found FA to be a good tool for uncovering responses to stress in natural populations, and we recommend exploring FA as a means of uncovering developmental instability in other systems that merit conservation interest.

Effects of repeated exposure to malathion on growth, food consumption, and locomotor performance of the western fence lizard ( Sceloporus occidentalis)

Effects of repeated pollutant exposure on growth, locomotor performance, and behavior have rarely been evaluated in reptiles. We administered three doses of malathion (2.0, 20, or 100 mg/kg body weight) to western fence lizards ( Sceloporus occidentalis) over an 81 day period. Eight and 23% mortality occurred at 20 and 100 mg/kg ( p = 0.079) and 85% of lizards in the 100 mg/kg group exhibited clinical symptoms of poisoning. Growth, food consumption, body condition index, and terrestrial locomotor performance were not significantly influenced by malathion. However, arboreal sprint velocity was significantly reduced in lizards receiving 100 mg/kg. Fifty percent of lizards in the 100 mg/kg group also refused to sprint in the arboreal setting ( p = 0.085). Based on these results, arboreal locomotor performance was the most sensitive metric of exposure we evaluated. Further study of compounds such as malathion is warranted due to highly variable application rates and exposure scenarios.

Clonal diversity of a lizard malaria parasite, Plasmodium mexicanum, in its vertebrate host, the western fence lizard: role of variation in transmission intensity over time and space

Within the vertebrate host, infections of a malaria parasite (Plasmodium) could include a single genotype of cells (single-clone infections) or two to several genotypes (multiclone infections). Clonal diversity of infection plays an important role in the biology of the parasite, including its life history, virulence, and transmission. We determined the clonal diversity of Plasmodium mexicanum, a lizard malaria parasite at a study region in northern California, using variable microsatellite markers, the first such study for any malaria parasite of lizards or birds (the most common hosts for Plasmodium species). Multiclonal infections are common (50-88% of infections among samples), and measures of genetic diversity for the metapopulation (expected heterozygosity, number of alleles per locus, allele length variation, and effective population size) all indicated a substantial overall genetic diversity. Comparing years with high prevalence (1996-1998 = 25-32% lizards infected), and years with low prevalence (2001-2005 = 6-12%) found fewer alleles in samples taken from the low-prevalence years, but no reduction in overall diversity (H = 0.64-0.90 among loci). In most cases, rare alleles appeared to be lost as prevalence declined. For sites chronically experiencing low transmission intensity (prevalence approximately 1%), overall diversity was also high (H = 0.79-0.91), but there were fewer multiclonal infections. Theory predicts an apparent excess in expected heterozygosity follows a genetic bottleneck. Evidence for such a distortion in genetic diversity was observed after the drop in parasite prevalence under the infinite alleles mutation model but not for the stepwise mutation model. The results are similar to those reported for the human malaria parasite, Plasmodium falciparum, worldwide, and support the conclusion that malaria parasites maintain high genetic diversity in host populations despite the potential for loss in alleles during the transmission cycle or during periods/locations when transmission intensity is low.

Assessment of lead uptake in reptilian prey species

As part of an investigation determining the trophically available fraction of metals in a model terrestrial food web, i.e., invertebrate prey to Western fence lizards (Sceloporus occidentalis), we evaluated the ability of several invertebrate prey to bioaccumulate lead and to form metals-rich granules, which are hypothesized to be non-available to predators. Crickets (Acheta domestica), tenebroid beetle larvae (Tenebrio molitor), and isopods (Porcellio scaber) were selected as model prey organisms. Lack of standard exposure methodologies for these species has presented a barrier to trophic transfer evaluations, as each species has particular requirements that create challenges for designing exposure conditions. We were able to devise exposure conditions for all three species that allow long-term exposure studies. All prey organisms accumulated lead from contaminated food, and for all species the majority of the accumulated Pb was associated with the exoskeleton (>50%), with metals-rich granules accounting for most of the remaining accumulated lead.

Selenomethionine Biotransformation and Incorporation into Proteins along a Simulated Terrestrial Food Chain

Selenium is an essential trace element in vertebrates, but there is a narrow concentration range between dietary requirement and toxicity threshold. Although a great deal is known about the biochemistry of Se from a nutritional perspective, considerably less attention has been focused on the specific biochemistry of Se as an environmental toxicant. Recent advances in hyphenated analytical techniques have provided the capability of quantifying specific chemical forms of Se in biological tissues as well as the distribution of Se among macromolecules. We applied liquid chromatography coupled to inductively coupled plasma mass spectrometryto investigate biotransformations of selenomethionine along a simulated terrestrial food chain consisting of selenomethionine exposed crickets (Acheta domesticus) fed to western fence lizards (Sceloporus occidentalis). Evidence was obtained for selenomethionine biotransformation as well as for sex-specific differences in the metabolism of Se compounds and their subsequent incorporation into proteins in the lizard. The results demonstrate the complexities involved in trophic transfer of Se due to the potential for extensive biotransformation and the species- and even sex-specific nature of these biotransformations.

EXPERIMENTAL TEST FOR PREMUNITION IN A LIZARD MALARIA PARASITE (PLASMODIUM MEXICANUM)

Premunition in Plasmodium spp. is the prevention of superinfection by novel genotypes entering an already established infection in a vertebrate host. Evidence for premunition was sought for the lizard malaria parasite, P. mexicanum, in its natural host, the fence lizard, Sceloporus occidentalis. Clonal diversity (= alleles for the haploid parasite) was determined with the use of 3 microsatellite markers. Both naturally infected lizards (N = 25) and previously noninfected lizards (N = 78) were inoculated intraperitoneally (IP) with blood from donor infections and followed over a 3-mo period. Compared to the success of clonal establishment in all the naive lizards (78/78 successful), clones entering preexisting infections had a significant disadvantage (9/25 successful). The number of preexisting clones (1-2 vs. 3-4) within recipient infections had no effect on the success of superinfection. Infections that excluded entering novel clones did not have higher initial asexual parasitemia, but had a higher initial density of gametocytes, suggesting they were older. Infections allowing superinfection experienced a higher final parasitemia.

Impaired terrestrial and arboreal locomotor performance in the western fence lizard ( Sceloporus occidentalis) after exposure to an AChE-inhibiting pesticide

We examined the effects of a commonly used AChE-inhibiting pesticide on terrestrial and arboreal sprint performance, important traits for predator avoidance and prey capture, in the western fence lizard ( Sceloporus occidentalis). Lizards were exposed to carbaryl (2.5, 25, and 250 μg/g) and were raced before and 4, 24, and 96 h after dosing. In the terrestrial setting, exposure to low concentrations of carbaryl had stimulatory effects on performance, but exposure to the highest concentration was inhibitory. No stimulatory effects of carbaryl were noted in the arboreal environment and performance in lizards was reduced after exposure to both the medium and highest dose of carbaryl. Our findings suggest that acute exposure to high concentrations of carbaryl can have important sublethal consequences on fitness-related traits in reptiles and that arboreal locomotor performance is a more sensitive indicator of AChE-inhibiting pesticide poisoning than terrestrial locomotor performance.

Ability of Transstadially Infected <I>Ixodes pacificus</I> (Acari: Ixodidae) to Transmit West Nile Virus to Song Sparrows or Western Fence Lizards

The hypothesis that Ixodes pacificus Cooley & Kohls (Acari: Ixodidae) may serve as a reservoir and vector of West Nile virus (family Flaviviridae, genus Flavivirus, WNV) in California was tested by determining the ability of this tick species to become infected with the NY99 strain of WNV while feeding on viremic song sparrows, to maintain the infection transstadially, and then to transmit WNV to recipient naive song sparrows and western fence lizards during the nymphal stage. The percentage of ticks testing positive by reverse transcription-polymerase chain reaction (RT-PCR) decreased from 77% of 35 larvae at day 6 after ticks were transferred to donor song sparrows (day of detachment) to 23% of 35 nymphs at 59 d postinfestation (approximately 19 d after molting to the nymphal stage). However, the percentage of ticks positive by RT-PCR from which infectious virus was recovered by Vero cell assay decreased from 59% on day 6 to 12% on day 59, even though there was no statistically significant decrease in the quantity of RNA within positive ticks. Attempts to improve the sensitivity of plaque assays by blind passage through C6/36 cell cultures were unsuccessful. These data indicated that ticks maintained viral RNA but not necessarily infectious virus over time. Nymphs from larvae that fed on song sparrows with peak viremias ranging from 7.2 to 8.5 log10 plaque-forming units (PFU) per ml were used in transmission attempts. From one to seven RNA-positive nymphal ticks engorged and detached from each of four recipient song sparrows or western fence lizards. Blood samples from sparrows and lizards remained negative, indicating that transmission did not occur. An additional four lizards inoculated with 1,500 PFU of WNV developed moderate viremias, ranging from 4.2 to 5.6 log10 PFU/ml. Our data and data from previous studies collectively indicated that ixodid ticks were not able to experimentally transmit WNV and therefore most likely would not be important vectors in WNV transmission cycles.

Western Bluebird Captures a Western Fence Lizard

I observed a male Western Bluebird (Sialia mexicana) capture a western fence lizard (Sceloporus occidentalis), the first reported incidence of vertebrate-directed raptorial behavior in the Western Bluebird. There are no previously published reports of Western Bluebirds capturing vertebrate prey, although there is one previous report of a Western Bluebird carrying an unidentified lizard in the manner of a prey item, and a few reports of predation on vertebrates by the congeneric Eastern Bluebird (Sialia sialis).

Energy acquisition and allocation in an ectothermic predator exposed to a common environmental stressor

Stressors are commonly encountered by organisms and often prove to be energetically costly. Certain stressors can simultaneously affect multiple components of an animal's energy budget and can either exacerbate energetic costs to the individual or offset one another. Here we used a commonly encountered stressor, the pesticide carbaryl, to examine the complex effects that acute environmental disturbances can have on energy expenditure, allocation, and acquisition, important processes that influence growth and reproduction. After exposing lizards ( Sceloporus occidentalis) to carbaryl, we measured their metabolism over a 48 h period and assessed their food consumption over 96 h. We found no difference in total energy expenditure among treatment groups, but lizards exposed to the highest dose of carbaryl allocated energy differently than other groups. Compared to controls, these lizards exhibited a 16–30% increase in standard metabolic rate (SMR), which was offset by a 45–58% decrease in additional energy expenditures. Lizards in the highest dose group also exhibited a 30–34% decrease in energy acquisition compared to controls. The net result was a 1.83 kJ decrease in energy assimilation, equivalent to 5 times their daily SMR requirements. Our results indicate that energetic consequences of stressors may result in complex energetic trade-offs, and emphasize the need to simultaneously examine the effect of stressors on multiple portions of an animal's energy budget.