Larval nematodes (Ascarops sp.) in stomach granulomas of the sagebrush lizard, Sceloporus graciosus

Wild populations of the sagebrush lizard (Sceloporus graciosus) are reported as paratenic hosts for third stage larvae of Ascarops sp. Larvae were found in stomach submucosal granulomas which averaged 350 microns in diameter and consisted of histiocytes and layers of fibrocytes. Prevalence of infection was 16% (27 of 170 infected); mean intensity was 3 (1 to 8 nematodes/host). Prevalence was significantly higher in male than female lizards.     

Malarial parasitism and male competition for mates in the western fence lizard,Sceloporus occidentalis

Summary The effect of malarial parasitism on the ability of male western fence lizards, Sceloporus occidentalis, to compete for access to females was assessed experimentally. Pairs of male lizards, one infected with the malarial parasite, Plasmodium mexicanum, and the other not infected, were matched by size and color and placed in large seminatural outdoor enclosures along with an adult female lizard. Infected males displayed to females and to other males less often than did noninfected male lizards. Noninfected lizards were dominant in social interactions more often than malarious animals, based on duration and intensity of agonistic encounters toward the other male, and time spent with the female. Thus, malarial infection hinders the ability of male fence lizards to compete for mates.     

Refractoriness of the western fence lizard (Sceloporus occidentalis) to the Lyme disease group spirochete Borrelia bissettii

The western fence lizard, Sceloporus occidentalis, is refractory to experimental infection with Borrelia burgdorferi sensu stricto, one of several Lyme disease spirochetes pathogenic for humans. Another member of the Lyme disease spirochete complex, Borrelia bissettii, is distributed widely throughout North America and a similar, if not identical, spirochete has been implicated as a human pathogen in southern Europe. To determine the susceptibility of S. occidentalis to B. bissettii, 6 na?ve lizards were exposed to the feeding activities of Ixodes pacificus nymphs experimentally infected with this spirochete. None of the lizards developed spirochetemias detectable by polymerase chain reaction for up to 8 wk post-tick feeding, infected nymphs apparently lost their B. bissettii infections within 1-2 wk after engorgement, and xenodiagnostic L. pacificus larvae that co-fed alongside infected nymphs did not acquire and maintain spirochetes. In contrast, 3 of 4 na?ve deer mice (Peromyscus maniculatus) exposed similarly to feeding by 1 or more B. bissettii-infected nymphs developed patent infections within 4 wk. These and previous findings suggest that the complement system of S. occidentalis typically destroys B. burgdorferi sensu lato spirochetes present in tissues of attached and feeding I. pacificus nymphs, thereby potentially reducing the probability of transmission of these bacteria to humans or other animals by the resultant adult ticks.     

Evolution of thermal physiology and growth rate between populations of the western fence lizard (Sceloporus occidentalis)

Summary Hatchling Sceloporus occidentalis from northern populations (central Oregon) grow more slowly than hatchlings from southern populations (southern California) in nature. In this study, I determine whether this difference in growth rate results from differences in thermal environment and/or in thermoregulatory behavior. To determine the degree to which the thermal environment affects growth rate among populations, I reared hatchings from the northern and southern populations in a cycling thermal regime in one of three experimental treatments differing in access to radiant heat (6, 9, or 12 h radiant heat; remainder of 24 h at 15°C). I also measured the body temperature that each individual voluntarily selected over the course of the daily activity cycle. Growth rate varied positively with duration of access to radiant heat. Within the three treatments, individual growth rate was positively correlated with body temperature. Moreover, the difference in growth rate between the northern and southern populations was due in part to differences in behavior — individuals from northern populations selected lower body temperatures. I found that significant variation in body temperature was associated with family membership, suggesting that thermal physiology has a genetic basis. Moreover, growth rate was correlated with body temperature among families in each population suggesting a genetic correlation underlies the phenotypic correlations. Thus, genetically based variation in thermal physiology contributes to differences in growth rate among individuals within a population as well as to differences among populations.     

Thyroxine affects behavioral thermoregulation but not growth rate among populations of the western fence lizard (Sceloporus occidentalis)

We investigated whether thyroxine influences hatchling growth rate of the western fence lizard (Sceloporus occidentalis) throught its effects on thermoregulatory behaviors. We reared control and thyroxine-injected hatchlings from three populations of S. occidentalis that differ in growth rate in a thermal gradient. We also measured the daily changes in body temperature and activity level (proportion of time spent out of retreat sites) of control and thyroxine-injected lizards. Previous studies have shown that within and among population differences in growth rate of the western fence lizard are correlated with the maintenance of high activity levels (proportion of time spent outside of retreat sites) and high body temperatures throughout the day. Growth rate was not influenced by injections of thyroxine. However, injections of thyroxine did elevate average daily body temperature and daily activity. Although administration of thyroxine uniformly increased the probability of activity throughout the day, it did not appear to alter the daily changes in activity. Previous studies have shown that the slower-growing hatchlings from northern populations exhibit a decline in activity during the later part of the thermal cycle, whereas the faster growing southern hatchlings maintain the same level of high activity throughout the thermal cycle. The decline in activity of northern populations was not prevented by thyroxine injection used in our current study. Northern lizards receiving exogenous thyroxine were still less active later in the day compared to early in the day, even though activity level throughout the day was elevated. Thus, the effects of thyroxine on temperature regulation observed in our study (general increase in activity level) appear to be unrelated to those aspects of temperature regulation (e.g., daily changes in behavioral thermoregulation) that are correlated with among population differences in growth rate. We also raised hatchlings in a cycling thermal regime (forced thermal cycle of 34°C:15°C, 12L:12D) where behavioral thermoregulation is not possible. The growth rate of lizards forced to cycle between 34°C:15°C on a daily basis was significantly lower than those lizards allowed to behaviorally thermoregulate, further underscoring the importance of the circadian pattern of thermoregulation for growth.Abbreviations GR growth rate - MR metabolic rate - SMR standard metabolic rate - SVL snout-vent length - T₄ thyroxine - T _b body temperature - T _e environmental temperature     

Inter-familiar differences in sprint speed of hatchling Sceloporus occidentalis (Reptilia: Iguanidae)

Hatchling Sceloporus occidentalis from the same dam had more similar sprint speeds than did hatchlings from different dams. Families (e.g. siblings from the same dam) of hatchlings differ significantly in body size. However, family differences in sprint speed remain significant when the size differences are removed statistically. Genetic differences among the families of lizards may account for most of the family differences in speed, although environmental or maternal effects may also contribute to family differences in sprint speed.     

Geographic variation in the effects of heat exposure on maximum sprint speed and Hsp70 expression in the western fence lizard Sceloporus occidentalis

We examined whether western fence lizards Sceloporus occidentalis occurring in thermally divergent environments display differential responses to high temperature in locomotor performance and heat-shock protein (Hsp) expression. We measured maximum sprint speed in S. occidentalis from four populations at paired latitudes and elevations before and after exposure to an experimental heat treatment and then quantified hind-limb muscle Hsp70 expression. Lizards collected from northern or high-elevation collection sites suffered a greater reduction in sprint speed after heat exposure than lizards collected from southern or low-elevation sites. In addition, lizards from northern collection sites also exhibited an increase in Hsp70 expression after heat exposure, whereas there was no effect of heat exposure on Hsp70 expression in lizards from southern collection sites. Across all groups, there was a negative relationship between Hsp70 expression and sprint speed after thermal stress. This result is significant because (a) it suggests that an increase in Hsp70 alone cannot compensate for the immediate negative effects of high-temperature exposure on sprint speed and (b) it demonstrates a novel correlation between an emergent property at the intersection of several physiological systems (locomotion) and a cellular response (Hsp70 expression). Ultimately, geographic variation in the effects of heat on sprint speed may translate into differential fitness and population viability during future increases in global air temperatures.     

Anonymous nuclear markers for the eastern fence lizard,Sceloporus undulatus

We present results from a screen for de novo variable nuclear loci using a genomic library approach in Sceloporus undulatus, the eastern fence lizard. We tested amplification success for 77 primer pairs in S. undulatus, Sceloporus occidentalis and Sceloporus grammicus. Many loci amplified in all three species suggesting that our primers will be useful for developing sequencing or single nucleotide polymorphism (SNP) genotyping markers in other sceloporine lizards. We also sequenced 19 loci, containing 158 variable sites, for 91 S. undulatus individuals. We report high levels of nucleotide variation in this species with an average of 38 SNPs per kilobase.     

Ascarops talpa sp. n. (Nematoda: Spirocercidae) from the formosan mole, Talpa micrura insularis, in Taiwan

Ascarops talpa sp. n. (Nematoda: Spirocercidae) is described from the small intestine of the Formosan mole, Talpa micrura insularis, from Taiwan. It is most similar to A. kutassi (Shul'ts, 1927) and A. tuvensis Sulimov, 1961. Ascarops kutassi differs by having a smaller ratio of left to right spicule length (1:4 vs. 1:5.5), no caudal alae, by its preanal papillae that are not equidistant from one another, and in its area rugosa which extends 1.25 mm anterior to the anus. Ascarops tuvensis differs by having a smaller ratio of left to right spicule length (1:3.8 vs. 1:5.5), by its preanal papillae that are not equidistant from one another, and in its area rugosa which covers only the ventral tip of the tail.     

Locomotor performance of hatchling fence lizards (Sceloporus occidentalis): Quantitative genetics and morphometric correlates

Summary We examined heritabilities and correlations among measures of locomotor performance (speed, stamina) and among possible morphometric determinants of performance (hindlimb span, tail length) in families of hatchling lizards (Sceloporus occidentalis). We were particularly interested in determining whether these traits were heritable and thus might potentially respond genetically to selection. Moreover, we wished to determine whether speed and stamina are negatively genetically correlated, as suggested bya priori physiological and empirical considerations. All four traits appeared to be significantly heritable. Broadsense heritabilities were 0.33–0.36 for speed, 0.35–0.36 for stamina, 0.45–0.51 for hindlimb span, and 0.46–0.47 for tail length. Contrary to expectations, speed and stamina were not negatively genetically correlated. Hindlimb span and tail length, however, were negatively genetically correlated (but not phenotypically correlated). Hindlimb span and stamina were positively phenotypically correlated. Thus, for example, selection for longer hindlimb span could potentially result in shorter tails, contrary to evolutionary predictions based only on phenotypic correlations.     

The influence of light on cone disk shedding in the lizard, Sceloporus occidentalis

The lizard, Sceloporus occidentalis has an all-cone retina. In lizards maintained on a 12-h light:12-h dark (12L:12D) cycle, a burst of cone outer segment (COS) shedding occurs 2 h after light offset (1400 h circadian time) (Young, R.W., 1977, J. Ultrastruct. Res. 61:172-72). In this investigation, we studied the effect of different lighting regimes on the pattern of cone disk shedding in this species. When lizards entrained to a 12L:12D cycle are kept in constant darkness (DD), the shedding peak is advanced approximately 2 h and the magnitude of shedding is reduced to 30% of control. COS increased in mean length from 12 micron in controls to 14 micron after one cycle in DD and maintained this length during a second cycle in DD. In constant light (LL), disk shedding was damped to approximately 10% of control values. Shedding synchrony in LL was also perturbed and therefore cyclic shedding bursts could not be distinguished. During LL there was a much larger increase in COS mean length than in DD. After one cycle of LL, COS length was 15 micron and after two cycles COS length exceeded 17 micron. When lizards entrained to 12L:12D are shifted to a 6L:18D regimen, the first shedding cycle is biphasic. The first peak of 5% shedding occurs 2 h after light offset whereas a second larger peak (13%) occurs according to the entrained schedule (1400 h). This manipulation separates out a dark-triggered and circadian shedding component, which is normally superimposed in lizards entrained to a 12L:12D cycle. When entrained lizards are placed in 36 h of LL followed by light offset, the peak shedding response after light offset is double the control response (53% vs. 27%). After 30 h of LL (lights off 90 degrees out of phase), there is a biphasic shedding response similar to the 6L:18D regimen although this time the dark-triggered shedding component is greater in magnitude then the circadian component. COS turnover is estimated by extrapolating from COS mean length increases during LL. From this method we obtained a 2.7-micron increase in COS length during each day in LL. If COS growth is not augmented during LL, this would yield a 4-5-d turnover time for the average 12.5-micron COS.     

Absence of measurable malaria-induced mortality in western fence lizards (Sceloporus occidentalis) in nature: a 4-year study of annual and over-winter mortality

Theoretical models of parasite virulence often quantify virulence by mortality. However, there is a lack of empirical studies of parasite-induced host mortality because it is often difficult to quantify in natural populations. I have estimated annual and over-winter mortality in a population of fence lizards (Sceloporus occidentalis) infected with a malaria parasite, Plasmodium mexicanum, in northern California. The duration of time a lizard was observed (an estimate of life-span) throughout the 4-year observation period, or following winter, was not related to either infection status or maximum parasitemia. In contrast to previous laboratory studies of this parasite-host system, I found no evidence of parasite-induced host mortality in nature.     

Seasonal alterations in adrenocortical cell function associated with stress-responsiveness and sex in the eastern fence lizard (Sceloporus undulatus)

We characterized steroidogenic properties of dispersed adrenocortical cells from field-active male and female eastern fence lizards (Sceloporus undulatus) to investigate whether alterations in cell function could, in part, explain seasonal variation in baseline and stress-induced plasma corticosterone (B). Lizards were collected during the breeding and postbreeding seasons and shortly prior to hibernation. Dispersed cells in vitro produced B, aldosterone (ALDO), and progesterone in response to 8-Br-cAMP, 25-(OH)cholesterol, adrenocorticotropin (ACTH; as little as 100 fM), and angiotensin II. Maximal progesterone, B, and ALDO responses to ACTH were roughly 1000%, 500%, and 100% greater than corresponding basal values. Angiotensin II was an effective steroidogenic stimulant but much less so than ACTH. Corticosteroid production exhibited considerable steroid-specific variation among seasons. Maximal ACTH-induced B production was lower in the postbreeding season than at either of the other two measurement points, essentially opposite to the pattern for ALDO. Males and females generally produced B at similar rates, but ALDO and progesterone showed numerous sex differences that usually covaried between the two steroids. Cellular sensitivity to 25-(OH)cholesterol and angiotensin II showed few sex differences or seasonal changes. In contrast, sensitivity to ACTH decreased markedly from the breeding to the postbreeding season in males, corresponding to the decrease in stress-responsiveness, and in both sexes was considerably lower prior to hibernation than during the breeding season. Under some conditions, plasma B may be limited by the production capacity of adrenocortical cells. In summary, seasonal variations in body condition, reproductive activity, and baseline and stress-induced plasma B may be attributed at least in part to alterations in adrenocortical cell steroidogenic function.     

Plasma androgens and their association with the reproductive cycle of the male fence lizard, Sceloporus undulatus

Seasonal variation in plasma androgen levels was determined for males of the lizard Sceloporus undulatus. Plasma androgens peaked in both spring and in fall prior to brumation, coinciding with periods of maximum testicular enlargement and advanced spermatogenic development. Plasma androgen concentrations were minimal during summer, following testicular regression and breeding in spring. Epididymal masses were positively correlated to plasma androgen levels, while abdominal fat body masses demonstrated a negative correlation.     

A second record of adult Ascarops strongylina (Rudolphi, 1819) (Nematoda: Spirocercidae) in a rodent host

Mature adults of Ascarops strongylina (Nematoda: Spirocercidae) were collected from a Manchurian zokor, Myospalax psilurus (Rodentia: Cricetidae), in Mongolia. This is the second record of adult A. strongylina from wild rodents and the first for M. psilurus as a definitive host of this nematode. Remnants of ground crustaceans (Porcellio sp.) and a dung beetle (Onthophagus sp.) were also found in the stomach of the zokor.     

Prevalence and abundance of Ixodes pacificus immatures (Acari: Ixodidae) infesting western fence lizards (Sceloporus occidentalis) in northern California: temporal trends and environmental correlates.

The prevalence and abundance of immature Ixodes pacificus ticks on western fence lizards (Sceloporus occidentalis) were examined in relation to time of year, host attributes (i.e., age, gender, and presence or absence of blood parasites), and 5 environmental characteristics, including topographic exposure and ground cover substrate, over a 2-year period in northern California. Lizards were infested with subadult ticks from early March until late July or early August, with peak median numbers of larvae and nymphs recorded in late April and early May of both years. Peak larval and nymphal abundances differed between years. The overall ratio of larvae to nymphs on adult male lizards was low, ranging from 0.80 in 1999 to 2.41 in 2000. Such intensive feeding of nymphs versus larvae on these lizards, which are reservoir-incompetent for Borrelia burgdorferi spirochetes, may explain previous observations of decreasing spirochetal infection prevalence from the nymphal to adult stage in northwestern California. Adult male lizards were more likely to be infested with nymphs and harbored greater abundances of larvae and nymphs than adult females. Lizards uninfected with blood parasites had more nymphs than infected lizards. The measured environmental characteristics could explain only a small percentage of the total variation observed in larval prevalence (22%) and in larval and nymphal abundance (12 and 3%, respectively).     

Chromosome number in the lizard genus Uta (family Iguanidae)

A chromosome number of 34 (12 macro- and 22 microchromosomes) was found to be characteristic of the bone marrow in 47 animals including males from the species Uta antiquus, and both males and females from the following species and subspecies: Uta stansburiana stansburiana, Uta stansburiana stejnegeri, Uta stansburiana elegans, Uta stansburiana klauberi, Uta stansburiana mannophorus, Uta nolascensis, Uta palmeri, and Uta squamata. — Diploid chromosome numbers of 34 and haploid numbers of 17 were found in the nine testis smears examined. — The presence of a large number of hypodiploid figures in the bone marrow smears is attributed to cell fragmentation and the problem of distinguishing the small microchromosomes. — Series of polyploid figures whose chromosome numbers increased in arithmetic rather than geometric progressions were observed in the testis dry smears. Possible alternatives for the origin of these figures are presented. — Problems encountered in the use of chromosome number as a taxonomic character are discussed.Supported in part by Research Grants GB-366 and GB-5416 from the National Science Foundation, and GM-15361 from the United States Public Health Service.