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Cost and benefits of lizard thermoregulation.

Lizards thermoregulate by behavioral and physiological adjustments. The resultant control over metabolic processes is generally assumed to be beneficial. However, these thermoregulatory adjustments have associated costs which, if extensive, make thermoregulation impractical. We extend this idea into an abstract mathematical, cost-benefit model of thermoregulation in lizards. Investigation of the model leads to a set of predictions which includes: (1) the physiologically optimal temperature is not always the ecologically optimal temperature; (2) thermoregulation is beneficial only when associated costs are low; (3) thermal specialists will normally thermoregulate more carefully than thermal generalists unless costs are high; and (4) lizards will thermoregulate more carefully if productivity of the habitat is increased or if exploitation competition is reduced. Data on lizards, where available, generally agree with these predicitions.

Animals↗

Ontogenetic scaling of bite force in lizards and turtles.

Because selection on juvenile life-history stages is likely strong, disproportionately high levels of performance (e.g., sprint speed, endurance, etc.) might be expected. Whereas this phenomenon has been demonstrated with respect to locomotor performance, data for feeding are scarce. Here, we investigate the relationships among body dimensions, head dimensions, and bite force during growth in lizards and turtles. We also investigate whether ontogenetic changes in bite performance are related to changes in diet. Our analyses show that, for turtles, head dimensions generally increase with negative allometry. For lizards, heads scale as expected for geometrically growing systems. Bite force generally increased isometrically with carapace length in turtles but showed significant positive allometry relative to body dimensions in lizards. However, both lizards and turtles display positive allometric scaling of bite force relative to some measures of head size throughout ontogeny, suggesting (1) strong selection for increased relative bite performance with increasing head size and (2) intrinsic changes in the geometry and/or mass of the jaw adductors during growth. Whereas our data generally do not provide strong evidence of compensation for lower absolute levels of performance, they do show strong links among morphology, bite force, and diet during growth.

Animals↗

Correlates of average daily metabolism of field-active zebra-tailed lizards (Callisaurus draconoides).

The extent of variation in reptile field metabolism, and its causal bases, are poorly understood. We studied the energetics of the insectivorous lizard Callisaurus draconoides at a site in the California Desert (Desert Center) and at a site at the southern tip of the Baja Peninsula (Cabo San Lucas; hereafter, Cabo). Reproducing Callisaurus were smaller at Cabo than at Desert Center. The allometry of metabolism with body mass can account for most differences in whole-animal metabolism. There was no significant effect of sex or source population on mass-adjusted metabolic rate in the laboratory (resting metabolism, measured by closed-system respirometry) or in the field (field metabolism, measured with doubly labeled water). The mass-adjusted resting metabolism and field metabolism of gravid females and the field metabolism of juvenile lizards were not significantly different from those of nonreproductive adults. Temperature had a significant effect on resting metabolism (Q10 = 2.7); fed lizards had resting metabolism that was 22% higher than that of fasted lizards; field metabolism was positively correlated with growth rate in juveniles; and field metabolism of adults increased from spring to late summer at Desert Center by 25%, probably because of longer activity period length and slightly higher activity period body temperature. We calculated from water influx and field metabolism that juveniles allocated 18% of their metabolizable energy intake to growth and that most energy deposited into eggs was transferred from energy stores rather than ingested in the weeks prior to laying.

Aging↗

Effects of testosterone on locomotor performance and growth in field-active northern fence lizards, Sceloporus undulatus hyacinthinus.

The role of steroids in locomotor performance and growth was examined in free-living lizards. Male northern fence lizards (Sceloporus undulatus hyacinthinus) with experimentally elevated plasma testosterone concentrations had greater sprint speed (+24%) and burst stamina (+17%) than sham-implanted males after 14-23 d in the field. This enhanced performance was associated with significant energetic costs, as the testosterone-implanted lizards had reduced growth rates, and, in a companion experiment, field-active testosterone-implanted lizards had smaller fat-body masses than controls after just 3-4 wk. These results suggest that, in addition to influencing a variety of behavioral and morphological traits, testosterone may play an important role in the regulation of locomotor performance. Also, natural levels of locomotor performance may be constrained, in part, by associated costs of elevated plasma testosterone concentrations.

Animals↗

Aponomma (Bothriocroton) glebopalma, n. subgen., n. sp., and Amblyomma glauerti n. sp. (Acari: Ixodida: Ixodidae), parasites of monitor lizards (Varanidae) in Australia.

Aponomma (Bothriocroton) glebopalma, n. subgen., n. sp., and Amblyomma glauerti, n. sp. (Acari: Ixodidae), are described from the monitor lizards Varanus glebopalma Mitchell and Varanus glauerti Mertens in Western Australia and the Northern Territory, Australia. The new subgenus is erected to accomodate A. (Bothriocroton) glebopalma, a new species with a deeply pitted and pilose scutum in both the adult and immature stages, unlike all other described species of Aponomma. This new species is found in an area where Aponomma fimbriatum Koch, a parasite of varanid lizards and various species of snakes, was the only known member of the genus found in both Western Australia and the Northern Territory. Amblyomma glauerti n. sp., an ornate Australian reptile tick found on the same two hosts in Western Australia and the Northern Territory, is also described. These two new species were recovered from formalin-preserved lizard specimens; therefore, nothing is known of their life cycles, except that all stages parasitize varanid lizards.

Animals↗

Chemoreception, symmetry and mate choice in lizards.

Research on fluctuating asymmetry (FA)-mediated sexual selection has focused almost exclusively on visual signals and ignored chemical communication despite the fact that many species rely on chemical signals for attracting mates. Female mate choice based on visual traits appears to be rare in lizards. However, the femoral glands of male lizards produce pheromones which might transmit chemical information about an individual's developmental stability. Therefore, we hypothesized that mate choice may be based on chemical cues. We analysed the effect of the developmental stability levels of males on the attractiveness of males' scents to females in a laboratory experiment with the lizard Lacerta monticola. When we offered two males of similar body size, females preferentially associated with the scents of males with low FA in their femoral pores and also with the scents of males with a higher number of femoral pores. This suggested that the females were able to discriminate the FA of the males by chemical signals alone and that the females preferred to be in areas marked by males of high quality, thus increasing their opportunities of mating with males of high quality. We suggest that the quality and/or amount of male pheromones could communicate the heritable genetic quality of a male to the female and thereby serve as the basis for adaptive female choice in lizards.

Animals↗

Lizard threat display handicaps endurance.

Honest-signalling theory asserts that threat displays reliably advertise attributes that influence fighting success. Endurance, as measured by treadmill performance, predicts the outcome of agonistic interactions among lizards. If threat displays in lizards function to advertise endurance capacity then variation in threat displays should correlate with endurance. I tested this prediction for the duration of threat posturing in male side-blotched lizards (Uta stansburiana) and examined whether threat displays act as quality handicaps, reliable signals that expend the attribute that is advertised. Individual variation in the duration of threat posturing correlated with endurance, while an experimental reduction of endurance diminished the duration of threat posturing. As expected of a quality handicap, endurance fell below baseline after display production. A restriction of aerobic metabolism can account for this effect. In threat posturing, lateral compression of the thorax may interfere with respiration or with circulation, limiting aerobic metabolism and causing a compensatory increase in anaerobic metabolism, thereby generating lactate and diminishing locomotor capacity. Concentrations of lactate measured after display production were higher than baseline, consistent with the proposed mechanism. By restricting aerobic metabolism, the threat posture can act as a quality handicap, simultaneously advertising and expending the endurance capacity of displaying lizards.

Aggression↗

Vitamin D supplementation increases the attractiveness of males' scent for female Iberian rock lizards.

Evolutionary theory proposes that signals used in sexual selection can only be stable if they are honest and condition dependent. However, despite the fact that chemical signals are used by many animals, empirical research has mainly focused on visual and acoustic signals. Vitamin D is an essential nutrient for lizards, but in some lizards its precursor (cholesta-5,7-dien-3-ol=provitamin D) is found in femoral gland secretions, which males use for scent marking and intraspecific communication. By allocating provitamin D to secretions, males might need to divert vitamin D from metabolism. This might be costly and condition dependent. We tested whether diet quality affected chemical signals of male Iberian rock lizards (Lacerta monticola) and its consequences for sexual selection. After experimental supplementation of dietary vitamin D, males increased the proportion of provitamin D in femoral secretions. Further experiments showed that females detected these changes in males' signals by chemosensory cues, and discriminated provitamin D, and changes in its concentration, from similar steroids (i.e. cholesterol) found in secretions. Moreover, females preferred areas scent marked by males with more provitamin D in their secretions. This mechanism would confer honesty to chemical signals of male lizards, and, thus, females may rely on it to select high-quality males. We suggest that the allocation of vitamins and other essential nutrients to either visual (e.g. carotenoids) or chemical ornaments might be the common basis of honest sexual displays in many animals.

Analysis of Variance↗

Evolutionary implications of phenotypic plasticity in the hindlimb of the lizard Anolis sagrei.

Species of Anolis lizards that use broad substrates have long legs, which provide enhanced maximal sprint speed, whereas species that use narrow surfaces have short legs, which permit careful movements. We raised hatchling A. sagrei in terraria provided with only broad or only narrow surfaces. At the end of the experiment, lizards in the broad treatment had relatively longer hindlimbs than lizards in the narrow treatment. These results indicate that not only is hindlimb length a plastic trait in these lizards, but that this plasticity leads to the production of phenotypes appropriate to particular environments. Comparison to hindlimb lengths of other Anolis species indicates that the range of plasticity is limited compared to the diversity shown throughout the anole radiation. Nonetheless, this plasticity potentially could have played an important role in the early stages of the Caribbean anole radiation.

Adaptation, Physiological↗

A comparative test of adaptive hypotheses for sexual size dimorphism in lizards.

It is commonly argued that sexual size dimorphism (SSD) in lizards has evolved in response to two primary, nonexclusive processes: (1) sexual selection for large male size, which confers an advantage in intrasexual mate competition (intrasexual selection hypothesis), and (2) natural selection for large female size, which confers a fecundity advantage (fecundity advantage hypothesis). However, outside of several well-studied lizard genera, the empirical support for these hypotheses has not been examined with appropriate phylogenetic control. We conducted a comparative phylogenetic analysis to test these hypotheses using literature data from 497 lizard populations representing 302 species and 18 families. As predicted by the intrasexual selection hypothesis, male aggression and territoriality are correlated with SSD, but evolutionary shifts in these categorical variables each explain less than 2% of the inferred evolutionary change in SSD. We found stronger correlations between SSD and continuous estimates of intrasexual selection such as male to female home range ratio and female home range size. These results are consistent with the criticism that categorical variables may obscure much of the actual variation in intrasexual selection intensity needed to explain patterns in SSD. In accordance with the fecundity advantage hypothesis, SSD is correlated with clutch size, reproductive frequency, and reproductive mode (but not fecundity slope, reduced major axis estimator of fecundity slope, length of reproductive season, or latitude). However, evolutionary shifts in clutch size explain less than 8% of the associated change in SSD, which also varies significantly in the absence of evolutionary shifts in reproductive frequency and mode. A multiple regression model retained territoriality and clutch size as significant predictors of SSD, but only 16% of the variation in SSD is explained using these variables. Intrasexual selection for large male size and fecundity selection for large female size have undoubtedly helped to shape patterns of SSD across lizards, but the comparative data at present provide only weak support for these hypotheses as general explanations for SSD in this group. Future work would benefit from the consideration of alternatives to these traditional evolutionary hypotheses, and the elucidation of proximate mechanisms influencing growth and SSD within populations.

Adaptation, Biological↗

Further studies on osmotic resistance of nucleated erythrocytes: observations with pigeon, peafowl, lizard and toad erythrocytes during changes in temperature and pH.

The osmotic resistance of pigeon, peafowl, lizard and toad erythrocytes at different temperatures and pH was studied. Erythrocytes from female pigeons showed greater osmotic resistance than those from males, but no sex difference appeared with erythrocytes from peafowls. Pigeon erythrocytes were more resistant and the red blood cell, packed cell volume and haemoglobin values were higher than those in peafowls. Although no significant differences appeared in their haematological values, erythrocytes from the lizard were more resistant than erythrocytes from the toad. At higher temperature, the osmotic resistance of pigeon, lizard and toad erythrocytes increased, while that of peafowl erythrocytes decreased. The resistance of toad erythrocytes decreased in acidic and alkaline solutions, but that of peafowl erythrocytes increased in both solutions. However, with pigeon and lizard erythrocytes, the resistance was unaltered in alkaline solution and decreased in acidic solution.

Animals↗

Isolation of a thermotolerant Paravahlkampfia sp. from lizard intestine: biology and molecular identification.

An amoeba was isolated from the intestines of several moribund pink-tongued skinks (lizards), Hemisphaeriodon gerrardi. Unusual features of this isolate were its ability to grow at temperatures of > or = 37 degrees C, and its inability to use Escherichia coli as a food source or to grow axenically on a variety of enriched culture media suitable for other soil amoeba isolates. Growth was abundant, however, on tissue culture cells, with amoebae clearing cell monolayers in approximately 48 h at 37 degrees C. Trophozoites had a vahlkampfiid-like morphology, moving by means of an anterior eruptive pseudopod. Cysts, round to slightly ovoid and lacking exit pores, were formed in culture. Tests for enflagellation of trophic amoebae were negative. Indirect immunofluorescence staining was negative for Naegleria fowleri and Willaertia sp. The isolate was sensitive to azithromycin, but not to amphotericin B, pentamidine isethionate, fluconazole, 5-fluorocytosine, and sulfadiazine. Phylogenetic analysis based on the PCR-amplified small subunit ribosomal DNA, identified the organism as Paravahlkampfia ustiana, an amoeba not previously isolated from either poikilothermic or homeothermic hosts. No evidence of pathology was seen in stained sections of lizard intestine, suggesting that the ameba was part of the normal fauna of the lizard gut. Its diet in the lizard intestine is unknown and the organism may have unusual growth requirements. Thus, P. ustiana joins other soil amoebae that have been isolated from mammals, amphibia, fish, and reptiles, which have the potential of becoming opportunistic pathogens.

Amoeba↗

Central cardiovascular effects of L-DOPA and clonidine in the lizard, Agama caudospinosa.

The effects of L-DOPA alone and in combination with two inhibitors of 1-aromatic-amino acid decarboxylase (DC), carbidopa and benserazid on the blood pressure (BP) and catecholamine (CA) content in the heart and brain has been investigated in the lizard Agama caudospinosa. The effect of clonidine on BP has also been examined in intact lizards and after spinal cord transection. It is concluded that dopamine (DA) is formed from L-DOPA in brain and heart and that carbidopa reduces the DA formation in the heart but penetrates the blood brain barrier (BBB) poorly, allowing a central decarboxylation of L-DOPA. Benserazid, on the other hand, penetrates the BBB and inhibits the central as well as the peripheral synthesis of DA from L-DOPA. L-DOPA in combination with carbidopa lowers the BP significantly in lizards tilted 45 degrees (head upwards) whereas the supine BP is unaffected. Benserazid pretreatment abolishes the decrease in BP after L-DOPA. L-DOPA alone does not produce any consistent changes in BP. Clonidine lowers BP in intact, tilted animals, while the BP is increased in spinal animals. The results indicate a central mode of action of L-DOPA as well as clonidine in the lizard studied.

Animals↗

Lizards infected with malaria: physiological and behavioral consequences.

In northern California, western fence lizards, Sceloporus occidentalis, are frequently parasitized by Plasmodium mexicanum, which causes malaria. Animals with this naturally occurring malarial infection are anemic: immature erythrocytes in peripheral blood become abundant (1 to 30 percent), and blood hemoglobin concentration decreases 25 percent. Maximal oxygen consumption decreases 15 percent and aerobic scope drops 29 percent in infected lizards; both correlate with blood hemoglobin concentration. Running stamina, but not burst running speed, is reduced in malarious lizards. There is a hierarchical relation between infection with malaria and effects on hematology, physiological function, and behavioral capacity. The results suggest that malarial infection may have significant effects on the ecology of lizard hosts.

Animals↗

Demonstration of prostaglandin activity in lungs of the rainbow lizard (Agama agama) by bioassay.

Extracts and perfusate effluents of lungs of the rainbow lizard (Agama agama) were assayed for prostaglandin-like activity. Results of differential bioassay and thin-layer chromatography suggested that the prostanoid was predominantly PGE2-like. The mean PGE2-like content of 10 lizard lung extracts was 2.9 micrograms g-1 wet weight compared with 146 ng g-1 in rat lungs. Mechanical pressure applied to the lung during perfusion through the pulmonary vasculature provoked the release of large quantities of PGE2-like material. This release was blocked by fatty acid cyclooxygenase inhibitors. Compared with guinea-pig and rat lungs, lizard lungs exhibited a markedly low capacity for inactivating PGE2. In view of an apparently high prostaglandin-forming and a low inactivating capacity, we speculate that under certain circumstances, lizard lungs may release vasoactive substances into the circulation.

Animals↗

Carbohydrate metabolism of the brain in the ageing male garden lizard: effect of cold shock on mitochondrial respiration and lactate/pyruvate ratio.

Mitochondrial respiration with succinate as substrate increased and the mitochondrial protein content declined in the brain of the ageing male garden lizard. The lactate/pyruvate ratio of both blood and brain did not show appreciable changes with advancing age. Cold-shock treatment induced a significant rise in the respiratory rate of brain mitochondria of young but not appreciably in that of middle-aged and old lizards. On the other hand both mitochondrial protein content and lactate/pyruvate ratio increased only in the brain of cold-shocked lizards of the old age-group. The lactate/pyruvate ratio in blood remained unaltered following cold-shock treatment of lizards of all three age-groups.

Aging↗

Cloning, sequencing, and expression of the amelogenin gene in two scincid lizards.

Our knowledge of the gene coding for amelogenin, the major enamel protein, is mainly based on mammalian sequences. Only two sequences are available in reptiles. To know whether the snake sequence is representative of the amelogenin condition in squamates, we have studied amelogenin in two scincid lizards. Lizard amelogenin possesses numerous conserved residues in the N- and C-terminal regions, but its central region is highly variable, even when compared with the snake sequence. This rapid evolution rate indicates that a single squamate sequence was not representative, and that comparative studies of reptilian amelogenins might be useful to detect the residues which are really important for amelogenin structure and function. Reptilian and mammalian enamel structure is roughly similar, but no data support amelogenin being similarly expressed during amelogenesis. By performing in situ hybridization using a specific probe, we showed that lizard ameloblasts express amelogenin as described during mammalian amelogenesis. However, we have not found amelogenin transcripts in odontoblasts. This indicates that full-length amelogenin is specific to enamel matrix, at least in this lizard.

Ameloblasts↗

Locomotion of lizards on inclines and perches: hindlimb kinematics of an arboreal specialist and a terrestrial generalist.

Arboreal animals, especially lizards, often traverse three-dimensional networks of narrow perches with variable and steep inclines, but the effects of both incline and narrow surfaces on the locomotor movement and function of limbs are poorly understood. Thus, we quantified the three-dimensional hindlimb kinematics of a specialized arboreal lizard, Chamaeleo calyptratus, moving horizontally, and up and down a 30 degrees incline on a narrow (2.4 cm) perch and a flat surface. We compared the flat-surface data of C. calyptratus with those of an anatomically generalized terrestrial lizard, Dipsosaurus dorsalis. Inclines had significant main effects for relatively few kinematic variables of C. calyptratus (11%) compared to D. dorsalis (73%). For C. calyptratus, the main effects of locomotor surface were nearly three times more widespread than those of incline. The foot of C. calyptratus was markedly anterior to the hip at footfall, primarily as a result of an unusually extended knee for a lizard. A large amount of knee flexion during early stance may be used by C. calyptratus to actively pull the body forward in a manner not found in D. dorsalis. Unexpectedly, the pelvic rotation of C. calyptratus greatly exceeded that of D. dorsalis and, unlike D. dorsalis, was not affected by incline. The more medial location of the foot of C. calyptratus on the narrow perch during stance was primarily a result of knee flexion rather than femur depression. Unlike previous qualitative descriptions of chameleons, our data for the hindlimb posture of C. calyptratus during stance indicate that the limb was not particularly erect.

Adaptation, Biological↗