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Clustering of related individuals in a population of the Australian lizard, Egernia frerei.

Stable social aggregations are rarely recorded in lizards, but have now been reported from several species in the Australian scincid genus Egernia. Most of those examples come from species using rock crevice refuges that are relatively easy to observe. But for many other Egernia species that occupy different habitats and are more secretive, it is hard to gather the observational data needed to deduce their social structure. Therefore, we used genotypes at six polymorphic microsatellite DNA loci of 229 individuals of Egernia frerei, trapped in 22 sampling sites over 3500 ha of eucalypt forest on Fraser Island, Australia. Each sampling site contained 15 trap locations in a 100 x 50 m grid. We estimated relatedness among pairs of individuals and found that relatedness was higher within than between sites. Relatedness of females within sites was higher than relatedness of males, and was higher than relatedness between males and females. Within sites we found that juvenile lizards were highly related to other juveniles and to adults trapped at the same location, or at adjacent locations, but relatedness decreased with increasing trap separation. We interpreted the results as suggesting high natal philopatry among juvenile lizards and adult females. This result is consistent with stable family group structure previously reported in rock dwelling Egernia species, and suggests that social behaviour in this genus is not habitat driven.

Animals↗

Mitochondrial DNA phylogeography of the Mesoamerican spiny-tailed lizards (Ctenosaura quinquecarinata complex): historical biogeography, species status and conservation.

Through the examination of past and present distributions of plants and animals, historical biogeographers have provided many insights on the dynamics of the massive organismal exchange between North and South America. However, relatively few phylogeographic studies have been attempted in the land bridge of Mesoamerica despite its importance to better understand the evolutionary forces influencing this biodiversity 'hotspot'. Here we use mitochondrial DNA sequence data from fresh samples and formalin-fixed museum specimens to investigate the genetic and biogeographic diversity of the threatened Mesoamerican spiny-tailed lizards of the Ctenosaura quinquecarinata complex. Species boundaries and their phylogeographic patterns are examined to better understand their disjunct distribution. Three monophyletic, allopatric lineages are established using mtDNA phylogenetic and nested clade analyses in (i) northern: México, (ii) central: Guatemala, El Salvador and Honduras, and (iii) southern: Nicaragua and Costa Rica. The average sequence divergence observed between lineages varied between 2.0% and 3.7% indicating that they do not represent a very recent split and the patterns of divergence support the recently established nomenclature of C. quinquecarinata, Ctenosaura flavidorsalis and Ctenosaura oaxacana. Considering the geological history of Mesoamerica and the observed phylogeographic patterns of these lizards, major evolutionary episodes of their radiation in Mesoamerica are postulated and are indicative of the regions' geological complexity. The implications of these findings for the historical biogeography, taxonomy and conservation of these lizards are discussed.

Animals↗

Omnivory in lacertid lizards: adaptive evolution or constraint?

Feeding specializations such as herbivory are an often cited example of convergent and adaptive evolution. However, some groups such as lizards appear constrained in the evolution of morphological specializations associated with specialized diets. Here we examine whether the inclusion of plant matter into the diet of omnivorous lacertid lizards has resulted in morphological specializations and whether these specializations reflect biomechanical compromises as expected if omnivores are constrained by functional trade-offs. We examined external head shape, skull shape, tooth structure, intestinal tract length and bite performance as previous studies have suggested correlations between the inclusion of plants into the diet and these traits. Our data show that omnivorous lacertid lizards possess modifications of these traits that allow them to successfully exploit plant material as a food source. Conversely, few indications of a compromise phenotype could be detected, suggesting that the evolution towards herbivory is only mildly constrained by functional trade-offs.

Adaptation, Physiological↗

Ontogenic sources of variation in sexual size dimorphism in a viviparous lizard.

To elucidate the developmental aspects of the evolution of sexual size dimorphism (SSD), an understanding of the sex-specific ontogeny of body size is critical. Here, we evaluate the relative importance of genetic and environmental determinants of SSD in juvenile common lizards (Lacerta vivipara). We examined the prenatal and post-natal effects of population density and habitat humidity on SSD, as well as the maternal effects of food availability, corticosterone level, humidity and heat regime during gestation. Analyses indicated strong prenatal and post-natal plasticity in body size per se and yielded three main results with respect to SSD. First, SSD in juvenile common lizards matches qualitatively the SSD observed in adults. Secondly, SSD was influenced by none of the prenatal factors investigated here, suggesting poor sex-biased maternal effects on offspring size. Thirdly, SSD was sensitive to post-natal habitat humidity, which positively affected growth rate more strongly in females than in males. Thus, natural variation in SSD in juvenile common lizards appears to be primarily determined by a combination of sex-biased genetic factors and post-natal conditions. We discuss the possibility that viviparity may constrain the evolution of sex-biased maternal effects on offspring size.

Animals↗

Chemical, immunological and biological properties of peptides like vasoactive-intestinal-peptide and peptide-histidine-isoleucinamide extracted from the venom of two lizards (Heloderma horridum and Heloderma suspectum).

Having previously isolated helodermin, the major peptide like vasoactive-intestinal-peptide and peptide-histidine-isoleucinamide, from the venom of the lizard Heloderma suspectum, we decided on a systematic exploration of all (VIP-PHI)-like peptides present in the venom of another lizard of the Helodermatidae family: Heloderma horridum. Six (VIP-PHI)-like peptides (PHH1 to 6) were purified to homogeneity from the venom of the lizard H. horridum with PHH3 and PHH4 representing two minor forms. All peptides cross-reacted in radioimmunoassays for helodermin and PHI but not for VIP. They yielded four fragments (T1 to T4) after trypsin digestion. T1, T2 and T3 showed the same retention time by reverse-phase HPLC and the same amino acid composition; the differences were confined to T4, the C-terminal sequence. PHH5 and PHH6 were found to be identical to synthetic helospectins I and II respectively. PHH1 and PHH3 probably resulted from a secondary modification of PHH5, while PHH2 and PHH4 derived from PHH6. Thus, the VIP-like peptides, previously called helospectins, are in fact typical of H. horridum venom. We confirmed that helodermin is the major (VIP-PHI)-like peptide of the venom of H. suspectum and observed its absence in H. horridum venom. Also, we found that positions 8 and 9 of helodermin are occupied by two Glu residues instead of two Gln as previously published. Helospectin-like material was also present in H. suspectum venom but in very small amount. In both venoms all VIP-like peptides were equally potent and efficient when tested for (a) their ability to occupy VIP as well as secretin receptors in rat pancreatic membranes and VIP receptors in rat liver membranes, and (b) the ensuing activation of adenylate cyclase in both membrane preparations.

Adenylyl Cyclases↗

Influence of forest fragmentation on community structure of frogs and lizards in northeastern Costa Rica.

To better understand responses of reptiles and amphibians to forest fragmentation in the lowland Neotropics, we examined community and population structure of frogs and lizards in the fragmented landscape surrounding La Selva Biological Station in the Sarapiquí region of northeastern Costa Rica. We used diurnal quadrats and nocturnal transects to sample frogs and lizards in nine forest fragments (1-7 ha each) and La Selva (1100 ha). Species richness in all fragments combined was 85% of that found in La Selva with comparable sampling effort. Richness varied from 10 to 24 species among forest fragments, compared with 36 species at La Selva. Lizard density was higher and frog density was lower in forest fragments than in La Selva. Community composition varied among sites and by fragment size class, and species occurrence was nested with respect to fragment area. Isolation and habitat variables did not significantly affect species richness, composition, or nestedness. We classified 34% of species as fragmentation sensitive because they were absent or occurred at low densities in fragments. Nevertheless, the relatively high diversity observed in the entire set of fragments indicates that preserving a network of small forest patches may be of considerable conservation value to the amphibians and reptiles of this region.

Animals↗

PCR-mediated recombination between Cryptosporidium spp. of lizards and snakes.

The presence or absence of genetic recombination has often been used as one of the criteria for Cryptosporidium species designation and population structure delineation. During a recent study of cryptosporidiosis in reptiles that were housed in the same room, 4 lizards were found to have concurrent infections of C. serpentis (a gastric parasite) and C. saurophilum (an intestinal parasite), and 6 snakes were concurrently infected with C. serpentis, C. saurophilum and a new Cryptosporidium as indicated by PCR-RFLP analysis of the SSU rRNA gene. DNA sequence analysis of cloned PCR products confirmed the diagnosis of mixed infections. Surprisingly, it appeared that 11 of the 22 clones (8 and 14 clones from a lizard and a snake, respectively) had chimeric sequences of two Cryptosporidium spp. BootScan analysis indicated the existence of recombinants among the cloned sequences and detection of the informative sites confirmed the BootScan results. Because the probability for genetic recombination between gastric and intestinal parasites is small, these hybrid sequences were likely results of PCR artifacts due to the presence of multiple templates. This was confirmed by PCR-sequencing analysis of single-copy templates using diluted DNA samples. Direct sequencing of 69 PCR products from 100- to 1,000-fold diluted DNAs from the same snake and lizard produced only sequences of C. serpentis, C. saurophilum and the unnamed Cryptosporidium sp. Thus, care should be taken to eliminate PCR artifacts when determining the presence of genetic recombination or interpreting results of population genetic studies.

Animals↗

Preliminary evidence for pineal-mediated extraretinal photoreception in relation to tail regeneration in the Gekkonid lizard, Hemidactylus flaviviridis.

The tail of the Gekkonid lizard Hemidactylus flaviviridis was autotomized and the animals were subjected to eight different photoperiodic schedules during the process of tail regeneration. Our previous observation had shown that long-day photoperiods stimulate the regeneration process, whereas short-day photoperiods depress it. Furthermore, it has also been demonstrated that the lateral eyes, or retinae, do not participate in photoperiodically significant photoreception in H. flaviviridis, as blinded Hemidactylus regenerated their autotomized tails like their sighted counterparts exposed to similar experimental photoregimes. In a further attempt to localize the site(s) of photoreception in these animals, one group of lizards had their heads painted with a mixture of Indian ink and Nile blue sulphate (II-NBS) [NL (HP)] in order to prevent light from penetrating to the pineal gland, and another group had their pineal glands surgically removed (pinealectomy, Px); the regenerative potentials were compared with their normal (NL) counterparts. Our results showed that the initiation and onset of regeneration, the daily growth rate, the total new growth (regenerate) produced at the end of regeneration and the total percentage replacement of the lost (autotomized) tails were significantly retarded in Px and NL (HP) animals, compared with the NL (unoperated and nonpainted) ones. Since pinealectomy as well as light deprivation to the pineal abolished the stimulatory influence of long-length photoperiods, the pineal gland is discussed here as a major transmitter of photic stimulus in lacertilian tail regeneration. It is presumed that in the lizard, as in mammals and some birds, the pineal gland acts by way of the neuroendocrine complex and/or the hypothalamohypophyseal axis.

Animals↗

Persistence of a plasma melatonin rhythm in constant darkness and its inhibition by constant light in the sleepy lizard, Tiliqua rugosa.

This study determined whether a blood plasma melatonin rhythm persists in constant photothermal environments in the sleepy lizard, Tiliqua rugosa. It builds upon an earlier investigation which provided equivocal results as to whether an in vivo melatonin rhythm persists in constant dark (DD) and light (LL) and temperature in this species. Using more frequent sampling points and new assay techniques, the present study showed that the melatonin rhythm persisted for at least 6 days at temperatures of 25 and 33 degrees C in constant dark (DD). The melatonin rhythm, however, was largely eliminated in constant light (LL) at 33 degrees C, thereby contradicting some previous findings in other species of reptiles where melatonin levels were apparently insensitive to an unexpected pulse of light at night. These results demonstrate that the sleepy lizard has a persistent, possibly circadian rhythm of melatonin in DD and constant temperature, and that the rhythm is inhibited by LL and constant temperature. Therefore, the sleepy lizard pineal gland may be an independent oscillator capable of driving the melatonin rhythm and be a transducer of the seasonally changing external photothermal environment.

Animals↗

Basilar-membrane motion in the alligator lizard: its relation to tonotopic organization and frequency selectivity.

In the alligator lizard the entire basilar membrane is accessible for measurements of its velocity by the Mössbauer method. Tests of the method indicate (1) the Mössbauer source can be placed on the basilar membrane without altering the signal-transmission properties of the cochlea, and (2) the source adheres to the basilar membrane. Isovelocity curves (IVCs) were constructed by plotting (as a function of tone frequency) the sound-pressure level at the tympanic membrane required to produce a specified velocity amplitude. IVCs from 21 lizards for source locations spanning the length of the basilar membrane indicate that basilar-membrane velocity does not vary systematically with longitudinal location as it does in mammalian cochleas. Measurements of velocity waveforms in two lizards do not indicate substantial nonlinearity in the inner-ear mechanical system. The frequency dependence of the basilar-membrane velocity is similar to that of the extrastapes velocity over the range 0.4 to 2 kHz. Thus, the tonotopic organization and frequency selectivity, which have been previously demonstrated in this species in responses of both auditory-nerve fibers and cells of the receptor organ, are apparently not primarily determined by basilar-membrane motion.

Animals↗

Parietal eyes in lizards: zoogeographical correlates.

Lizards without parietal eyes tend to be restricted to low latitudes, whereas lizards with parietal eyes are successful at higher latitudes also. These zoogeographical data, along with current theories of parietal eye physiology, strongly suggest that the overall significance of the parietal eye to lizards as a group is that it facilitates survival at higher latitudes, thus making possible the exploitation of a wider variety of habitats.

Adaptation, Biological↗

Gene dosage at the lactate dehydrogenase b locus in triploid and diploid teiid lizards.

Triploid parthenogenetic lizards which are heterozygous at the lactate dehydrogenase b locus show approximately two doses of the b allele and approximately one dose of the b' allele. This finding agrees with the triploid karyotype composed of two haploid complements of chromosomes indistinguishable from those of species carrying the b allele and one complement indistinguishable from that of the species which carries the b' allele. Densitometry of electrophoretic patterns of lactate dehydrogenase from heart muscle and from kidney indicates approximately equal expression of each of the b alleles in the triploid lizards. Heterozygouis diploid lizards from the same parthenogenetic species, in agreement with their karyotype, show equal doses of b and b' alleles.

Alleles↗

Enterobacteriaceae isolated from iguanid lizards of west-central Texas.

The prevalence of members of the family Enterobacteriaceae in the intestines of seven species of iguanid lizards native to west-central Texas was determined. Of the 67 lizard specimens examined, 48.7% were infected with Salmonella and 9% were infected with Salmonella arizonae. Two lizard species (Sceloporus olivaceus and Crotaphytus collaris) were shown to have a 100% prevalence of Salmonella.

Animals↗

Assessment and conservation of the giant gomeran lizard (Gallotia bravoana).

In June 1999, a species of lizard previously considered extinct was rediscovered on the island of La Gomera, Spain. The giant gomeran lizard, Gallotia bravoana, is highly endangered and at risk of extinction by the feral cat population. A conservation effort to save the species was initiatived by the regional and central Spanish government and six lizards were captured and brought into captivity. This paper describes the results of the initial assessment of the health of the animals, by a physical examination and by haematological, biochemical, radiographical and coprological investigations. The methods used to maintain the animals in captivity are described and the first captive breeding success is reported.

Animal Diseases↗

Comparative haematology and blood chemistry of endangered lizards (Gallotia species) in the Canary Islands.

Blood samples were taken from the ventral coccygeal vein of 15 El Hierro giant lizards (Gallotia simonyi) (seven females and eight males), six La Gomera giant lizards (Gallotia bravoana) (four males and two females) and four Tenerife giant lizards (Gallotia intermedia) (two males and two females), and 31 blood parameters were measured. Among the haematological parameters there were significant differences between the three species in heterophils, azurophils and lymphocytes, but no significant differences in red blood cell count, white blood cell count, haemoglobin, packed-cell volume, monocytes, eosinophils and basophils. In terms of blood chemistry there were significant differences between the three species in cholesterol, triglycerides, glucose, sodium, chloride, urea, uric acid, total proteins, prealbumin, albumin and gamma globulins, but no significant differences in calcium, potassium, aspartate aminotransferase, alanine aminotransferase, creatine kinase, bile acids, alpha-1 and alpha-2 globulins and beta globulins.

Animals↗

Molecular evolution of satellite DNA repeats and speciation of lizards of the genus Darevskia (Sauria: Lacertidae).

Satellite DNA repeats were studied in Caucasian populations of 18 rock lizard species of the genus Darevskia. Four subfamilies (Caucasian Lacerta satellites (CLsat)I-IV) were identified, which shared 70%-75% sequence similarity. The distribution of CLsat subfamilies among the species was studied. All the species could be divided into at least 3 clades, depending on the content of CLsat subfamilies in each genome: "saxicola", "rudis", and "mixta" lizards. CLsatI was found in all studied species, but in very different quantities; the "saxicola" group contained this subfamily predominantly. The "rudis" group also contained CLsatIII, and the "mixta" group carried considerable amounts of CLsatII. The highest concentrations of CLsatI and CLsatII were detected in 2 ground lizards--D. derjugini and D. praticola, respectively. D. parvula predominantly carried CLsatIII. CLsatIV was found only in the Crimean species D. lindholmi. The distribution patterns of satellite subfamilies show possible postglacial speciation within the genus Darevskia. A hybrid origin of species that possess 2 or 3 CLsat subfamilies and important clarifications to the systematics of the genus are proposed.

Animals↗

Thermocyclic entrainment of lizard blood plasma melatonin rhythms in constant and cyclic photic environments.

We assessed how chronic exposure to 6-h cryophase temperatures of 15 degrees C in an otherwise 33 degrees C environment entrains the rhythm of blood plasma melatonin rhythms in lizards (Tiliqua rugosa) subjected to constant dark (DD), constant light (LL), and to 12:12-h light-dark cycles (12L:12D). The peak of the melatonin rhythm was entrained by the cryophase temperature of the thermocycle in DD and LL, irrespective of the time at which the cryophase temperature was applied. Comparable thermocycles of 6 h at 15 degrees C imposed on a 12L:12D photocycle, however, affected the amplitude and phase of the melatonin rhythm, depending on the phase relationship between light and temperature. Cold pulses in the early light period and at midday resulted, respectively, either in low amplitude or nonexistent melatonin rhythms, whereas those centered in or around the dark phase elicited rhythms of high amplitude. Supplementary experiments in 12L:12D using two intermittent 6-h 15 degrees C cryophases, one delivered in the midscotophase and another in the midphotophase, elicited melatonin rhythms comparable to those in lizards subjected to constant 33 degrees C and 12L:12D. In contrast, lizards subjected to 12L:12D and a 33 degrees C:15 degrees C thermocycle, whose thermophase was aligned with the photophase, produced a threefold increase in the amplitude of the melatonin rhythm. Taken together, these results support the notion that there is an interaction between the external light and temperature cycle and a circadian clock in determining melatonin rhythms in Tiliqua rugosa.

Acclimatization↗

Control of pulmonary surfactant secretion from type II pneumocytes isolated from the lizard Pogona vitticeps.

Pulmonary surfactant, a mixture consisting of lipids and proteins and secreted by type II cells, functions to reduce the surface tension of the fluid lining of the lung, and thereby decreases the work of breathing. In mammals, surfactant secretion appears to be influenced primarily by the sympathetic nervous system and changes in ventilatory pattern. The parasympathetic nervous system is not believed to affect surfactant secretion in mammals. Very little is known about the factors that control surfactant secretion in nonmammalian vertebrates. Here, a new methodology for the isolation and culture of type II pneumocytes from the lizard Pogona vitticeps is presented. We examined the effects of the major autonomic neurotransmitters, epinephrine (Epi) and ACh, on total phospholipid (PL), disaturated PL (DSP), and cholesterol (Chol) secretion. At 37 degrees C, only Epi stimulated secretion of total PL and DSP from primary cultures of lizard type II cells, and secretion was blocked by the beta-adrenoreceptor antagonist propranolol. Neither of the agonists affected Chol secretion. At 18 degrees C, Epi and ACh both stimulated DSP and PL secretion but not Chol secretion. The secretion of surfactant Chol does not appear to be under autonomic control. It appears that the secretion of surfactant PL is predominantly controlled by the autonomic nervous system in lizards. The sympathetic nervous system may control surfactant secretion at high temperatures, whereas the parasympathetic nervous system may predominate at lower body temperatures, stimulating surfactant secretion without elevating metabolic rate.

Acetylcholine↗