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Head shape and bite performance in xenosaurid lizards.

Bite performance in lizards influences many aspects of the animal's lifestyle. During feeding, inter- and intrasexual interactions, and defensive behavior, the ability to bite hard might be advantageous. Although biomechanical considerations predict clear relations between head shape and bite performance, this has rarely been tested. Here we investigate the effect of head shape on bite performance in three closely related species of xenosaurid lizards. Our data show that in this family of lizards, bite performance is mainly determined by head height, with high headed animals biting harder than flat headed ones. Species clearly differ in head shape and bite performance and show a marked sexual dimorphism. The dimorphism in head shape also results in an intersexual difference in bite performance. As head height is the major determinant of bite performance in xenosaurid lizards, trade offs between a crevice dwelling life-style and bite performance seem to occur. The evolutionary implications of these results are discussed. J. Exp. Zool. 290:101-107, 2001.

Animals↗

Fat body ovarian relationship in the garden lizard, Calotes versicolor (Daud.).

Abdominal fat body mass of Calotes versicolor showed annual changes that were universal related to the changes in ovarian somatic (GSI) and hepatosomatic (HSI) indices. Fat bodies were absent in late breeding phase (June-August). Thirty day fatectomy (FBX) during prebreeding phase significantly reduced GSI, HSI, and total number of extrastromal follicles; also, recruitment of vitellogenic follicles was arrested and ateretic follicles increased. The FBX during postbreeding phase had no such effect, whereas in 30 day ovariectomised (OvX) lizards in prebreeding phase fat body mass significantly increased but HSI decreased. However, in lizards in prebreeding phase, E2 caused a significant decrease in fat body mass and an increase in HSI, while during the postbreeding phase there was a significant increase in HSI but the fat bodies were not affected. The above findings suggest that the development of the first clutch of vitellogenic follicles in the lizard utilises lipids stored in the fat bodies and that the growth of the subsequent clutches of vitellogenic follicles is met through the intake of food, which is abundant in the latter part of the breeding phase. The fat bodies are not needed for the growth of previtellogenic follicles. The fact that lipolytic action of E2 occurs only during the breeding phase suggests that responsiveness of the fat bodies to the steroid is related to the reproductive phase and that during postbreeding phase of the lizard they become refractory to E2.

Animals↗

Temporal expression of thyroid hormone receptor alpha1 in the liver of the lizard Podarcis sicula.

The effects of thyroid hormones on metabolism and development are mediated by thyroid hormone receptors (TRs). To gain a better understanding of the potential role of thyroid hormone receptors in the liver of the lizard Podarcis sicula, we have evaluated the expression of TRs during the more critical periods of the annual variations of thyroid activity. The results obtained have indicated that in the liver of the lizard P. sicula there are three transcripts: mRNA of 5.0 kb for TRalpha1, mRNA of 2.6 kb for TRalpha2, and 6.0 kb band, which represent unprocessed heteronuclear RNA, encoding unspliced primary transcripts of RNA prior to their processing into the mature TRalpha1 and TRalpha2. By means of slot-blot, we are able to determine that there is a change in the expression of TRs that occurs in the liver during the annual cycle of thyroid activity. A major expression registers in May, when the lizard thyroid gland shows the maximal activity. The combination of molecular biology with immunohistochemistry revealed that hepatic cells were also TRalpha IR positive. Particularly intense immunostaining was present in the cell nuclei of animals sacrified in May. These observations suggest that in lizard P. sicula the thyroid hormone (T3) might regulate hepatic activity, modulating TR mRNA levels.

Analysis of Variance↗

Loricrin-like immunoreactivity during keratinization in lizard epidermis.

Two modalities of keratinization are present in lizard epidermis: alpha (soft-pliable corneous layers) and beta (hard and inflexible corneous layers). While beta-keratinization is probably due to the synthesis of a new (beta)-keratin gene product, alpha keratinization resembles in part that of mammalian epidermis. The goal of this study was to test whether a sulfur-rich molecule similar to the mammalian corneous cell envelope protein loricrin is also present in lizard epidermis. This was done using X-ray microanalysis and immunocytochemical and ultrastructural methods. In the epidermis of the lizard Podarcis muralis small (0.1-0.3 microm) to large (1-5 microm) keratohyalin-like granules (KHLGs) are produced in alpha-keratinizing cells, especially in the clear layer. Small KHLGs contain sulfur and show weak filaggrin-like and stronger loricrin-like immunoreactivities. The latter is also present in keratinizing alpha-layers but is absent in the beta layers. Large KHLGs in the clear layer derive from the aggregation of the small granules with other components, including lipid material. These large granules show some loricrin-like immunoreactivity and contain sulfur and phosphorous, histidine, but not filaggrin-like immunoreactivity. It is suggested here that phosphorous derives from their phospholipid component. The present study shows that the modality of alpha-keratinization of lizard epidermis resembles that of mammals and suggests that the basic molecular mechanisms of keratin aggregation and formation of the corneous cell envelope were already present in the therapsid line of reptiles from which mammals evolved.

Animals↗

Cytological evaluation of spermatogenesis within the germinal epithelium of the male European wall lizard, Podarcis muralis.

The annual cytological changes to the male germinal epithelium were investigated in an introduced population of European wall lizards (Podarcis muralis). Testicular tissues were collected, embedded, sectioned by an ultramicrotome, and stained with the PAS procedure followed by a toluidine counterstain. Spermatogenesis in the lizard is divided into the proliferative, meiotic, and maturational phases. Wall lizards have a prenuptial pattern of spermatogenesis, where sperm development begins immediately prior to and continues through the months of breeding (April-June). The testis then involutes, undergoes a short period of quiescence, and recrudescence commences in mid-July. Germ cells undergo proliferation, meiosis, and the early stages of spermiogenesis (maturation) from late July through December. However, the late stages of spermiogenesis are retarded from December through February. Spermiogenesis continues at an accelerated pace from March through May, leading to a single massive spermiation event through the month of June. Although spatial relationships are seen between germ cells within the seminiferous epithelium, accumulation of spermatids during winter and acceleration of elongation in spring prevents determination of consistent cellular associations between early and late developing germ cells within the wall lizard testis. This temporal germ cell development is different from the consistent spatial development seen within seasonally breeding birds and mammals and may represent an evolutionary intermediate in terms of amniotic germ cell development.

Animals↗

Limb and tail lengths in relation to substrate usage in Tropidurus lizards.

A close relationship between morphology and habitat is well documented for anoline lizards. To test the generality of this relationship in lizards, snout-vent, tail, and limb lengths of 18 species of Tropidurus (Tropiduridae) were measured and comparisons made between body proportions and substrate usage. Phylogenetic analysis of covariance by computer simulation suggests that the three species inhabiting sandy soils have relatively longer feet than do other species. Phylogenetic ANCOVA also demonstrates that the three species inhabiting tree canopies and locomoting on small branches have short tails and hind limbs. These three species constitute a single subclade within the overall Tropidurus phylogeny and analyses with independent contrasts indicate that divergence in relative tail and hind limb length has been rapid since they split from their sister clade. Being restricted to a single subclade, the difference in body proportions could logically be interpreted as either an adaptation to the clade's lifestyle or simply a nonadaptive synapomorphy for this lineage. Nevertheless, previous comparative studies of another clade of lizards (Anolis) as well as experimental studies of Sceloporus lizards sprinting on rods of different diameters support the adaptive interpretation.

Animals↗

Ultrastructural evidence for secretion from the epithelium of ampulla ductus deferentis of the fan-throated lizard Sitana ponticeriana Cuvier.

Among reptiles, an ampulla ductus deferentis has been reported only in Squamata. Fairly detailed studies are available only for two species, the lizard Calotes versicolor (Fam: Agamidae) and the snake Seminatrix pygaea (Fam: Colubridae). The light microscopic study on C. versicolor revealed the ampulla to be a prominent organ, whereas the light and transmission electron microscopic study in S. pygaea revealed it to be discernable only in histological preparations. Further, the epithelium of the ductal portion of vas deferens as well as the ampulla of C. versicolor appears to contribute to the seminal plasma and can also phagocytose dead sperm, whereas in S. pygaea neither of these roles has been established. Thus, we hypothesize that there may be variations in the anatomy, histology, and the role of the vas deferens in general, and the ampulla in particular, of the squamate reptiles. In this study, the ductus deferens of the small fan-throated lizard Sitana ponticeriana (Fam: Agamidae) was subjected to light and transmission electron microscopic analysis. In this lizard the ampulla is more prominent than in C. versicolor. The epithelium of the ductal portion of vas deferens consists of principal cells (with features reflecting roles in endocytosis and phagocytosis of dead sperm), dark cells (which are absent in the epithelium of the ductal portion of vas deferens of snakes), and basal cells. The ampulla of S. ponticeriana is differentiated into storage and glandular portions. The epithelium of the storage portion is like that in the ductal portion of the vas deferens, whereas that of the glandular portion, consisting of dark and light principal cells and foamy cells, is tall and forms into smooth villous folds. All three cell types show evidence for a role in secretion, in all likelihood different from each other, for release into the lumen to contribute to seminal plasma. These cells do not provide evidence of a role in phagocytosis of dead sperm. It appears that within the Squamata, the ductal ampulla differs in structure as well as function. We suggest that the ductal ampulla of agamid lizards is a composite gland of the ampulla ductus deferentis and seminal vesicles of mammals.

Animals↗

In vitro differentiation of tooth buds from embryos and adult lizards (L. gravenhorsti): an ultrastructural comparison.

It is well established that the capacity for teeth to differentiate "in vitro" depends upon: a) the age of the embryonic rudiments at the time of excision and b) the number of cells within each tissue type which are capable of differentiating into organ culture. This paper studies ultrastructural aspects of tooth buds grown in vitro from lizard embryos and compares these characteristics with those observed in dental germs grown in situ in older lizard embryos. Moreover, we report the self-differentiation in vitro dental tissues from adult lizard and compare this phenomenon with the main features of a morphogenetic field. Our results suggest that approximately in the first third of gestation in L. gravenhorsti the dental buds has already acquired the capacity for self-differentiation in vitro. The ultrastructural observations show that there are no significant differences between odontoblasts and ameloblasts in situ and in vitro. The tooth from "adult lizards," isolated by combined microsurgical and enzymatic procedure and cultured in semisolid-liquid medium were also able to differentiate teeth. This phenomenon implies that self-differentiation is not rigidly determined, and that in these animals the tooth tissues represents a continuous morphogenetic field throughout the animal's life. This property is intrinsic, resides in the isolated tooth tissues, and is relatively independent of external factors. In addition, these studies indicate that the chick chorio-allantoic membrane and the semisolid-liquid culture medium supply the majority of the factors required for development of these tissues.

Ameloblasts↗

Testis and epididymis of the Indian wall lizard (Hemidactylus flaviviridis): effects of flutamide on FSH and testosterone influenced spermatogenesis, Leydig cell, and epididymis.

To determine the separate spermatogenic actions of FSH and testosterone, adult male lizards Hemidactylus flaviviridis with recrudescent testes were administered the non-steroidal antiandrogen flutamide either alone or in combination with FSH or testosterone, and the histology and histochemistry of the testes and ductus epididymides were studied. Flutamide-treated animals displayed a marked hypertrophy of Leydig cells. A few spermatids were also seen in testis of more than half the animals treated with flutamide. Flutamide also produced a significant increase of primary spermatocytes; no spermatids were observed in controls. A significant inhibition of spermatogenesis was noted in lizards treated either with testosterone alone or in combination with flutamide. Ovine FSH treatment caused a significant stimulation of spermatogenesis, as indicated by the increase of primary and secondary spermatocytes and the transformation of secondary spermatocytes into spermatids or, in a few cases, into spermatozoa. A considerable depletion of sudanophilic lipid and moderate delta 5-3 beta-hydroxysteroid dehydrogenase activity was noted in the Leydig cells of FSH-treated animals indicating enhanced steroidogenesis. Similar results were obtained when lizards were treated with flutamide + FSH. The effects of simultaneous treatment of flutamide with FSH or testosterone on ductus epididymidis revealed that flutamide markedly inhibited the epithelial cell height and lumen diameter with a loss of luminal content when compared to FSH or testosterone-treated lizards.

Animals↗

Relationship between adenylate cyclase sensitivity to follitropin and FSH receptor mRNA expression in the ovary of the lizard Podarcis sicula.

In mammals, gonadal functions are regulated by two pituitary gonadotropins, follicle-stimulating hormone (FSH) and luteinizing hormone (LH), that interact with gonadal membrane receptors to activate adenylate cyclase. In comparison to mammalian systems, in squamate reptiles a reduced amount of information exists on gonadotropins and their related receptors. This study is aimed at clarifying if, in the lizard Podarcis sicula, the ovarian sensitivity to FSH is correlated to the reproductive cycle and to the expression of membrane receptors involved in the hormone recognition. The results demonstrate that the ovarian adenylate cyclase responsiveness to FSH parallels ovarian functions, being maximal during the ovulatory period. The ovarian sensitivity to FSH is also related to oocyte growth and vitellogenesis. Northern blot analyses reveal that the FSH receptor mRNA is maximally expressed in vitellogenic oocytes during the reproductive period. These results suggest that, in lizard ovary, hormone activation of adenylate cyclase is mediated by de novo synthesis of receptors specifically involved in FSH recognition. In lizards treated in vivo with FSH during the pre-ovulatory period, adenylate cyclase becomes refractory to further FSH stimulation 2 hr after treatment, but sensitivity to the hormone is restored after 2 weeks. Nevertheless, while the restored level of activity never exceeds that observed during the nonreproductive period, the expression level of FSH receptor mRNAs is significantly enhanced in these animals. These results suggest that in lizard the processes that regulate ovarian growth, vitellogenesis, and ovulation are controlled by a complex network of signals including gonadotropin, FSH receptor expression, and adenylate cyclase.

Adenylyl Cyclases↗

Regeneration of retinal axons in the lizard Gallotia galloti is not linked to generation of new retinal ganglion cells.

Using anterograde tracing with HRP and antibodies (ABs) against neurofilaments, we show that regrowth of retinal ganglion cell (RGC) axons in the lizard Gallotia galloti commences only 2 months after optic nerve transection (ONS) and continues over at least 9 months. This is unusually long when compared to RGC axon regeneration in fish or amphibians. Following ONS, lizard RGCs up-regulate the immediate early gene C-JUN for 9 months or longer, indicating their reactive state. In keeping with the in vivo data, axon outgrowth from lizard retinal explants is increased above control levels from 6 weeks, reaches its maximum as late as 3 months, and remains elevated for at least 1 year after ONS. By means of BrdU incorporation assays and antiproliferating cell nuclear antigen immunohistochemistry, we show that the late axon outgrowth is not derived from new RGCs that might have arisen in reaction to ONS: no labeled cells were detected in lizard retinas at 0.5, 1, 1.5, 3, 6, and 12 months after ONS. Conversely, numbers of RGCs undergoing apoptosis were too low to be detectable in TUNEL assays at any time after ONS. These results demonstrate that retinal axon regeneration in G. galloti is due to axon regrowth from the resident population of RGCs, which remain in a reactive state over an extended time interval. Neurogenesis does not appear to be involved in RGC axon regrowth in G. galloti.

Animals↗

S100 immunoreactive glial cells in the forebrain and midbrain of the lizard Gallotia galloti during ontogeny.

We identified S100 immunoreactive cells in the brain of the lizard Gallotia galloti during ontogeny using immunohistochemical techniques for light and electron microscopy. In double labeling experiments with antibodies specific for S100A1 and S100B (anti-S100) and proliferative cell nuclear antigen (anti-PCNA), myelin basic protein (anti-MBP), phosphorylated neurofilaments (SMI-31), glial fibrillary acidic protein (anti-GFAP), or glutamine synthetase (anti-GS), we detected S100-like immunoreactivity in glial cells but never in neurons. Restricted areas of the ventricular zone were stained in the hypothalamus from E32 to postnatal stages, and in the telencephalon at E35, E36, and in adults. S100 immunoreactivity was observed predominantly in scattered PCNA-negative cells that increased in number from E35 to the adult stage in the myelinated tracts of the brain and had the appearance of oligodendrocytes. Quantitative analysis revealed that all of the S100-positive glial cells were GFAP-negative, whereas most of the S100-positive glial cells were GS-positive. Ultrastructurally, most of these S100-positive/GS-positive glial cells resembled oligodendrocytes of light and medium electron density. In adult lizards, a small subpopulation of astrocyte-like cells was also stained in the pretectum. We conclude that in the lizard S100 can be considered a marker of a subpopulation of oligodendrocytes rather than of astrocytes, as is the case in mammals. The S100-positive subpopulation of oligodendrocytes in the lizard could represent cells actively involved in the process of myelination during development and in the maintenance of myelin sheaths in the adult.

Animals↗

Responsiveness of adenylate cyclase to pituitary gonadotropins and evidence of a hormone-induced desensitization in the lizard ovary.

Gonadotropins (FSH and LH) affect several mammalian gonadal functions. In particular, FSH stimulates oogonial proliferation and oocyte growth, while LH regulates ovulation and progesterone secretion. In lacertilian reptiles, gonadal function is also regulated by pituitary gonadotropins, but which hormone controls ovarian activities and the mechanisms of action are unknown. The present study aimed to clarify mechanisms of action of pituitary gonadotropins on the ovary of Podarcis sicula (Lacertilia). The data demonstrate that mammalian gonadotropins FSH and LH produce a threefold stimulation of adenylate cyclase activity in follicular membranes, while hCG and TSH are less effective, causing a twofold increase in adenylate cyclase activity. Neurotransmitters such as dopamine, serotonin, and catecholamines have no effect on enzyme activity. The action of mammalian FSH and LH on the ovary mimics the effect of homologous hormones: in lizard ovaries incubated in vitro in the presence of isolated homologous pituitary glands, the intracellular cAMP level increased by 50% with respect to control ovaries. Mammalian gonadotropins appear homologous to lizard gonadotropin(s): Southern blot analyses show that the lizard genome contains nucleotide sequences homologous to those encoding for mammalian beta FSH and beta LH. Both homologous and heterologous desensitization of adenylate cyclase activity occurs in the lizard ovary. In fact, responsiveness of adenylate cyclase to gonadotropin stimulation is abolished in animals 2 hr after in vivo treatment with FSH. Sensitivity to gonadotropin stimulation is restored 2 weeks after the beginning of the in vivo treatment. Desensitization was also observed in ovaries incubated in vitro with mammalian FSH or with isolated pituitary glands.

Adenylyl Cyclases↗

Number of mast cells in the Harderian gland of the lizard Podarcis sicula sicula (Raf): the annual cycle and its relation to environmental factors and estradiol administration.

The Harderian gland of the lizard Podarcis sicula sicula (Raf) contains connective tissue type mast cells whose numbers vary during the year showing two peaks, one in spring the other in winter. No sex differences are found throughout the year. Thermal and photoperiodic manipulations indicate that only temperature influences mast cell number (MCN) both in winter and in summer but not in spring. In animals exposed to high temperatures in February (but not in May) MCN declined, while exposure to low temperature in July had the opposite effect. Estradiol treatment of the February and April lizards increased MCN, an effect counteracted by the synthetic antiestrogen tamoxifen; in July lizards, this did not occur. In animals exposed to a high temperature in February, estradiol had no effect, as in animals exposed to low temperatures in July. These data suggest that in spring MCN seems to be more responsive to hormonal stimuli rather than external cues (temperature), while in summer MCN is more sensitive to temperature than to hormonal stimuli (estradiol). Both humoral and external factors are concluded to influence mast cell numbers in the Harderian gland of the lizard P. sicula sicula.

Animals↗

Hepatic outer-ring deiodinase in a Mexican endemic lizard (Sceloporus grammicus).

The kinetic characterization of the outer-ring deiodination pathway using rT(3) (rT(3)-ORD) in male, female, and pregnant female livers of an endemic lizard, Sceloporus grammicus, is reported. The ORD pathway does not have the characteristics of deiodinase type II; it is exclusively carried out by deiodinase type I (DI). DI enzymatic activity in lizard liver contains one of the highest activities reported in vertebrates. This activity is sexually dimorphic, with males presenting the highest activity during the reproductive season. The properties of this enzyme correspond to those described in mammals, such as specificity for rT(3), susceptibility to inhibition by 6-n-propyl-2-thiouracil and gold-thioglucose, cofactor requirement, and kinetic pattern. Unlike other vertebrates, the lizard DI exhibits conspicuous stability in the thermal range of 15 to 42 degrees C and in the pH range of 5.0 to 9.0. Male true kinetic constants exhibit a direct correlation with temperature. This is in agreement with short-term adaptation to microenvironmental changes and the feasible expression of enzymatic forms/variants which, together, endow this lizard species with a greater adaptation to natural daily ambient thermal fluctuations.

Animals↗

Effects of corticosterone on activity and home-range size of free-ranging male lizards.

The purpose of our study was to examine whether corticosterone (B) affects the spacing behavior of free-ranging male side-blotched lizards (Uta stansburiana). Furthermore, we wanted to determine if the density, reflected in seasonal changes in population size, or behavior, as a result of hormonal manipulation, of "neighboring" males influences these effects. Field studies were conducted on four naturally isolated "neighborhoods" of lizards. Half of the males on three of the sites were randomly implanted with either saline or B, while on the fourth site all males were implanted with B. Pre- and postimplant home-ranges and activity levels were determined. Home-range size and activity level were significantly reduced by corticosterone if normally aggressive saline-implanted males were also present in the neighborhood. However, B had no effect on home-range if all males in the neighborhood were implanted with B. Space lost by B-implanted males was incorporated into the home-ranges of saline-implanted males so that the sum of all the male home-ranges in a neighborhood remained unchanged after implantation. These results suggest that elevated B levels put male lizards at a competitive disadvantage and, therefore, force these lizards to reduce their home-range.

Animals↗

Phylogeny of xantusiid lizards: concern for data and analysis.

Phylogenetic analyses of new DNA sequence data from the mitochondrial 16S rRNA gene in xantusiid lizards support the intergeneric relationships obtained previously (S.B. Hedges, R.L. Bezy, and L.B. Maxson, 1991, Mol. Biol. Evol. 8:767-780) with data from the 12S rRNA and cytochrome b (cyt b) genes. The total data set now includes 1028 alignable sites, 471 of which are variable and informative for the distance analyses and 281 of which are informative for the parsimony analyses. Crother and Presch (1993), Mol. Phylogenet. Evol. 1:289-294) claim that their reanalyses of our 12S rRNA and cyt b sequence data do not support a robust phylogeny for xantusiid lizards. However, that conclusion is not supported by their own analyses of the combined data from those two genes, which result in the same phylogenetic tree of xantusiid genera that we obtained in the original study with the same method (maximum parsimony). This result was unchanged when Crother and Presch eliminated sites containing insertions/deletions and ambiguities, and when transversions were weighted. The less robust result for the separate cyt b analyses, probably due to the smaller size of the data set, was already noted (Hedges et al., 1991). We believe that the best estimate of relationships, in this case, is obtained by combining the sequence data from these tightly linked mitochondrial genes. We also refute the criticisms by Crother and Presch of the neighbor-joining method. To correct for a higher rate of transitions in mitochondrial sequence data, they weight transversions more heavily (5x) than transitions. We present theoretical criticisms of this weighting method and advocate the use of scaled corrections available with distance methods. Crother and Presch also claim that a robust phylogeny of xantusiid lizards is not obtained when some morphological data (13 informative characters) are combined with the molecular data in a single analysis. However, there are serious problems with their morphological data and methods of analysis. We reevaluate the three pivotal morphological characters in their alternative phylogeny for xantusiid genera and demonstrate that none of the three provides unambiguous support for their alternative arrangement. Crother and Presch implement a new approach whereby the entire morphological data set is weighted equally to the molecular data set in a combined analysis, thus resulting in a very inflated weight assigned to each character in the small morphological data set. Using this rationale, each of the 13 informative morphological characters would receive a greater than million-fold weight if combined with the sequence data for the entire genome of these lizards.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

Molecular phylogenetics, tRNA evolution, and historical biogeography in anguid lizards and related taxonomic families.

Phylogenetic relationships among lizards of the families Anguidae, Anniellidae, Xenosauridae, and Shinisauridae are investigated using 2001 aligned bases of mitochondrial DNA sequence from the genes encoding ND1 (subunit one of NADH dehydrogenase), tRNA(Ile), tRNA(Gln), tRNA(Met), ND2, tRNA(Trp), tRNA(Ala), tRNA(Asn), tRNA(Cys), tRNA(Tyr), and COI (subunit I of cytochrome c oxidase), plus the origin for light-strand replication (O(L)) between the tRNA(Asn) and the tRNA(Cys) genes. The aligned sequences contain 1013 phylogenetically informative characters. A well-resolved phylogenetic hypothesis is obtained. Because monophyly of the family Xenosauridae (Shinisaurus and Xenosaurus) is statistically rejected, we recommend placing Shinisaurus in a separate family, the Shinisauridae. The family Anniellidae and the anguid subfamilies Gerrhonotinae and Anguinae each form monophyletic groups receiving statistical support. The Diploglossinae*, which appears monophyletic, is retained as a metataxon (denoted with an asterisk) because its monophyly is statistically neither supported nor rejected. The family Anguidae appears monophyletic in analyses of the DNA sequence data, and statistical support for its monophyly is provided by reanalysis of previously published allozymic data. Anguid lizards appear to have had a northern origin in Laurasia. Taxa currently located on Gondwanan plates arrived there by dispersal from the north in two separate events, one from the West Indies to South America and another from a Laurasian plate to Morocco. Because basal anguine lineages are located in western Eurasia and Morocco, formation of the Atlantic Ocean (late Eocene) is implicated in the separation of the Anguinae from its North American sister taxon, the Gerrhonotinae. Subsequent dispersal of anguine lizards to East Asia and North America appears to have followed the Oligocene drying of the Turgai Sea. The alternative hypothesis, that anguine lizards originated in North America and dispersed to Asia via the Bering land bridge with subsequent colonization of Europe and Morocco, requires a phylogenetic tree seven steps longer than the most parsimonious hypothesis. North African, European, and West Asian anguines were isolated from others by the rapid uplift of Tibet in the late Oligocene to Miocene. Phylogenetic analysis of evolutionary changes in the gene encoding tRNA(Cys) suggests gradual reduction of dihydrouridine (D) stems by successive deletion of bases in some lineages. This evolutionary pattern contrasts with the one observed for parallel elimination of the D-stem in mitochondrial tRNAs of eight other reptile groups, in which replication slippage produces direct repeats. An unusual, enlarged TpsiC (T) stem is inferred for tRNA(Cys) in most species.

Animals↗