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Ontogeny of somatostatin immunoreactive neurons in the medial cerebral cortex and other cortical areas of the lizard Podarcis hispanica.

The ontogeny of somatostatin immunoreactive interneurons in the cerebral cortex of the lizard Podarcis hispanica has been studied in histological series of embryos, perinatal specimens, and adults. Somatostatin immunoreactive interneurons appear in the early stages of lizard cerebral cortex ontogeny, their number increases during embryonary development, reaches a peak in early postnatal life, and decreases in adult lizards. The first somatostatin immunoreactive somata in the lizard forebrain appeared on E36, and they were located in non cortical areas. Then, on E39 and later, somatostatin immunoreactive neurons were seen in the lizard cortex in a rostral-to-caudal spatial gradient, which parallels that of the normal histogenesis of the lizard cerebral cortex. On E39, labelled somata were seen in the medial and dorsal cortex inner plexiform layers; immunoreactive puncta and dendritic processes were detectable in the inner plexiform layer of the medial cortex. On E40, labelled neurons were observed in the inner plexiform layer of the lateral cortex; labelled processes were found in the inner plexiform layers (dorsomedial, dorsal, and lateral cortices) and the outer plexiform layers (medial and dorsomedial cortices). At hatching (P0), some somatostatin immunoreactive neurons populated the external plexiform layer of the dorsomedial cortex. On P28, groups of labelled neurons appeared in the cell layer of dorsal and lateral cortices, reaching the adult-mature pattern of somatostatin immunoreactivity in the lizard cerebral cortex, i.e., labelled somata and dendritic processes populating the inner plexiform layers in addition to an axonic labelled plexus in the outermost part of the outer plexiform layers. Immunoreactive somata and processes occupied all the cortical areas, but they were especially abundant in the dorsomedial cortex. Proliferating Cell Nuclear Antigen (PCNA) immunostaining in the same histological series revealed that the number of PCNA immunoreactive nuclei in the subjacent proliferative neuroepithelium followed an inverse-complementary evolution to somatostatin, suggesting some temporal relationship between somatostatin immunoreactive cells and neurogenesis in the lizard cerebral cortex.

Animals↗

Food chemical cues elicit general and population-specific effects on lingual and biting behaviors in the lacertid lizard Podarcis lilfordi.

Actively foraging lizards are capable of identifying prey using only chemical cues sampled by tongue-flicking, and the relatively few omnivorous and herbivorous lizards tested similarly can identify both animal and plant foods from chemical cues. Whether lizards that eat plants respond to cues specific to preferred plant types and whether there is geographic variability in responses to cues from various plants correlated with the importance of those plants in local diets is unknown. In three populations of an omnivorous lacertid, the Balearic lizard Podarcis lilfordi, we studied chemosensory sampling and feeding responses to chemical cues from plant and animal foods presented on cotton swabs. Each lizard population is endemic to one islet off the coast of Menorca, Balearic Islands, Spain. Lizards in all three populations discriminated chemical cues from plant and animal foods from control substances. Our results extend findings of prey chemical discrimination and plant chemical discrimination in omnivores, increasing confidence that correlated evolution has occurred between plant diet and chemosensory response to palatable plants. There were no consistent differences among populations in tongue-flicking and biting responses to stimuli from flowers of syntopic and allopatric plant species. The lizards may respond to cues indicative of palatability in a wide range of plant species rather than exhibiting strong responses only to locally available plant species. Nevertheless, tongue-flicking and biting frequencies varied among plant species, perhaps indicating food preferences. In addition, there were differences among populations in tongue-flick rates, latency to bite, and licking behavior. Licking was observed in only one lizard population as a response to floral chemicals from only one of the plants species tested, raising the possibility of a population-specific linkage between identification of a particular plant species and performance of an appropriate feeding response.

Animals↗

Saurian malaria in Kenya: epidemiological features of malarial infections in lizard populations of the West Pokot District.

During investigations into the prevalence of malarial parasites among lizards in the West Pokot District in Kenya, 179 lizards comprising eight species were caught. Examination of the Giemsa-stained smears made from their blood showed that 34 lizards were infected with Plasmodium species. Fifteen lizards were infected with a single species of Plasmodium and 19 carried multiple infections, the maximum, in four lizards, was four species. There were 19 combinations of parasite infections. Seventeen Plasmodium species were identified, the commonest being P. icipeensis. Only two of the eight lizard species were infected: the skink Mabuya striata and the agamid Agama agama. Eight of the Plasmodium species infected both; another eight species infected M. striata only but three of these have been described from different lizard families elsewhere in Africa. P. robinsoni infected A. agama only, although it was first described from another lizard family in another part of Africa. The epidemiological significance of these results is discussed.

Animals↗

Age-related differences in the response of hepatic oxygen consumption to thermal stress in the male garden lizard.

The main aim of the present study is to compare the thermal adaptation capacity of young, middle-aged and old lizards with respect to hepatic oxygen consumption. Cold exposure (1 h at 0-4 degrees C) caused a decline in endogenous oxygen consumption in young but did not evoke significant changes in middle-aged and old lizards. On the other hand, with 5 mM succinate the rate of respiration increased marginally only in cold-exposed old lizards. This suggests that the metabolic response to cold is age-dependent in these lizards. Exposure of homogenates at 40 +/- 1 degree C for 15 min led to a rise in the hepatic oxygen consumption of young but not in that of either middle-aged or old lizards. There were no changes in the rate of oxygen consumption upon exposure for a similar period at 45 +/- 1 degree C of liver homogenates from lizards of three different age groups. However, exposure at 51 +/- 1 degree C resulted in a higher degree of decline in the rate of hepatic oxygen consumption in middle-aged and old than in young lizards. Thus, the young lizards appear to be better-adapted to hyperthermic stress than the middle-aged and old.

Adaptation, Physiological↗

Resting metabolic expenditure as a potential source of variation in growth rates of the sagebrush lizard.

Along an elevational gradient on SW Utah, sagebrush lizards (Sceloporus graciosus) exhibit an unexpected pattern of growth. Lizards from a high elevation population grow faster than lizards from two populations at lower elevations despite shorter daily and seasonal activity. Results from a common environment study of growth suggest that the differences in growth are not due to adaptation to local environmental conditions. In this study, I test the hypothesis that higher growth rates in lizards from high elevation may be attributable to reduced resting metabolic expenditure compared to that of lizards from populations at two lower elevations. Resting metabolic rates were measured for individuals from each of the study populations across different times of day and over a broad range of temperatures. Under the same laboratory conditions, field-caught lizards from the high elevation population exhibited lower metabolic rates when compared to lizards from lower elevations. Daily resting metabolic expenditures were calculated using the observed metabolic rates coupled with estimates of daily activity. Daily resting metabolic expenditure was 50% greater for individuals from the two lower elevation populations, which could result in 12.5% more energy that could be potentially allocated to growth for lizards from high elevation. Such energetic savings may be able to explain differences in the patterns of growth observed in nature.

Acclimatization↗

Thermal dependence of chemical assessment of predation risk affects the ability of wall lizards, Podarcis muralis, to avoid unsafe refuges.

Prey often respond to predators by increasing refuge use, but some refuges may expose prey to other types of predators. Because predators are not always visible inside refuges, their chemical stimuli may provide early warning of their presence. However, in ectotherms, chemosensory detection of predators may be thermally dependent. Lizards often hide in cold refuges, where their body temperature (Tb) may decrease, and this might affect their ability to detect chemicals of saurophagous snakes that ambush inside refuges. We examined the effects of Tb of wall lizards, Podarcis muralis, on their ability to detect chemicals of smooth snakes, Coronella austriaca. Differential tongue flick (TF) rates and behavioral patterns of lizards in response to scent of smooth snakes showed that lizards with optimal Tb discriminated and avoided the snake's scent, whereas lizards with suboptimal Tb did not. We also examined the importance of this effect on movement patterns and refuge use by lizards in terraria with safe (odorless) or unsafe refuges (snake-scented). While results of this experiment were variable, there was evidence of snake avoidance in that lizards at optimal Tb spent less time in snake-scented refuges relative to odorless refuges. Therefore, this study provides evidence that chemosensory discrimination of snake chemicals is thermally dependent, and, thus, suboptimal Tb impedes a lizard's ability to avoid snake-scented refuges.

Analysis of Variance↗

Lizards as hosts for immature Ixodes scapularis (Acari: Ixodidae) in North Carolina.

Previously archived museum specimens of lizards collected throughout North Carolina were examined for Ixodes scapularis (Say). Lizards (n = 1,349) collected in 80 of North Carolina's 100 counties were examined. Lizards with ticks were collected in 23 (29%) of the 80 counties from which lizards were examined. I. scapularis was detected on 8.7% (n = 117) of the lizards and was the sole species of tick obtained from lizards. Immature ticks were most frequently found on the southeastern five-lined skink, Eumeces inexpectatus, and the eastern glass lizard, Ophisaurus ventralis. Larvae were most frequently found on the six-lined racerunner, Cnemidophorus sexlineatus. One C. sexlineatus harbored 177 larvae and 2 nymphs. Nymphs were most frequently observed on E. inexpectatus. The majority of counties (chi 2, P < 0.01) where ticks were found on lizards were in the Coastal Plain.

Animals↗

Effects of constant lightness, darkness and parachlorophenylalanine treatment on tail regeneration in the lizard Ophisops elegans macrodactylus: macroscopic, biochemical and histological changes.

A better understanding of regenerative growth is very important for the development of new potential strategies. Recently, the pineal secretory product melatonin was shown to stimulate the regeneration process. In this study, we carried out an experimental investigation of tail regeneration in young adult lizards, Ophisops elegans macrodactylus Berthold, 1842, addressing the role of melatonin on the regeneration process. Lizards were divided into three groups: constant light-exposed control group (n = 13), constant dark-exposed group (n = 15) and parachlorophenylalanine (p-CPA) treatment group exposed to constant light (n = 15). Using a tail autotomy procedure, the effects of p-CPA treatment on connective tissue together with neural tissue and vascular tissue in regenerating tail in lizards were investigated. p-CPA (400 microg/kg body mass) was injected from day 0 to 30 days after autotomy. p-CPA treatment produced a significant increase in the length of the regenerated tail compared with light-exposed control and dark-exposed lizards. Total collagen content was found to be higher in p-CPA-treated animals in comparison with other groups. Histologically, a higher percentage of connective tissue and vascular tissue and a lower percentage of neural tissue were found in the regenerated tails of the p-CPA-treated lizards. Importantly, the percentage of neural tissue in lizards in the dark-exposed animals was higher than that in animals of both light-exposed and p-CPA treatment groups. Thus, it is clear that p-CPA has a stimulatory influence on fibroblast collagen production and vascularization of the regenerated tail in the lizards. Furthermore, it seems that the neural regeneration process was markedly enhanced in lizards exposed to continuous darkness. Based on the results of our study, it is suggested that melatonin may be an active factor that speeds up the rate of tail regeneration in lacertilians.

Animals↗

Molecular identification and analysis of Borrelia burgdorferi sensu lato in lizards in the southeastern United States.

Lyme borreliosis (LB) group spirochetes, collectively known as Borrelia burgdorferi sensu lato, are distributed worldwide. Wild rodents are acknowledged as the most important reservoir hosts. Ixodes scapularis is the primary vector of B. burgdorferi sensu lato in the eastern United States, and in the southeastern United States, the larvae and nymphs mostly parasitize certain species of lizards. The primary aim of the present study was to determine whether wild lizards in the southeastern United States are naturally infected with Lyme borreliae. Blood samples obtained from lizards in Florida and South Carolina were tested for the presence of LB spirochetes primarily by using B. burgdorferi sensu lato-specific PCR assays that amplify portions of the flagellin (flaB), outer surface protein A (ospA), and 66-kDa protein (p66) genes. Attempts to isolate spirochetes from a small number of PCR-positive lizards failed. However, PCR amplification and sequence analysis of partial flaB, ospA, and p66 gene fragments confirmed numerous strains of B. burgdorferi sensu lato, including Borrelia andersonii, Borrelia bissettii, and B. burgdorferi sensu stricto, in blood from lizards from both states. B. burgdorferi sensu lato DNA was identified in 86 of 160 (54%) lizards representing nine species and six genera. The high infection prevalence and broad distribution of infection among different lizard species at different sites and at different times of the year suggest that LB spirochetes are established in lizards in the southeastern United States.

Animals↗

Paramyxovirus infection in caiman lizards (Draecena guianensis).

Three separate epidemics occurred in caiman lizards (Dracaena guianensis) that were imported into the USA from Peru in late 1998 and early 1999. Histologic evaluation of tissues from necropsied lizards demonstrated a proliferative pneumonia. Electron microscopic examination of lung tissue revealed a virus that was consistent with members of the family Paramyxoviridae. Using a rabbit polyclonal antibody against an isolate of ophidian (snake) paramyxovirus, an immunoperoxidase staining technique demonstrated immunoreactivity within pulmonary epithelial cells of 1 lizard. Homogenates of lung, brain, liver, or kidney from affected lizards were placed in flasks containing monolayers of either terrapene heart cells or viper heart cells. Five to 10 days later, syncytial cells formed. When Vero cells were inoculated with supernatant of infected terrapene heart cells, similar syncytial cells developed. Electron microscopic evaluation of infected terrapene heart cells revealed intracytoplasmic inclusions consisting of nucleocapsid strands. Using negative-staining electron microscopy, abundant filamentous nucleocapsid material with a herringbone structure typical of the Paramyxoviridae was observed in culture medium of infected viper heart cells. Seven months following the initial epizootic, blood samples were collected from surviving group 1 lizards, and a hemagglutination inhibition assay was performed to determine presence of specific antibody against the caiman lizard isolate. Of the 17 lizards sampled, 7 had titers of < or =1:20 and 10 had titers of >1:20 and < or =1:80. This report is only the second of a paramyxovirus identified in a lizard and is the first to snow the relationship between histologic and ultrastructural findings and virus isolation.

Animals↗

Muscle fiber-type variation in lizards (Squamata) and phylogenetic reconstruction of hypothesized ancestral states.

Previously, we found that phrynosomatid lizards, a diverse group common in the southwestern USA, vary markedly in fiber-type composition of the iliofibularis (a hindlimb muscle important in locomotion). Phrynosomatidae comprises three subclades: the closely related sand and horned lizards, and their relatives the Sceloporus group. The variation in muscle fiber-type composition for 11 phrynosomatid species is attributable mainly to differences between the sand- and horned-lizard subclades. Here, we expand the phrynosomatid database with three additional species and compare these results with data collected for 10 outgroup (distantly related) species. Our goal was to determine if the patterns found in Phrynosomatidae hold across a broader phylogenetic range of the extant lizards and to elucidate the evolution of muscle fiber-type composition and related traits. To allow for meaningful comparisons, data were collected from species that are primarily terrestrial and relatively small in size (3.5-65 g body mass). Results indicate that the fiber-type variation observed within the Phrynosomatidae almost spans the range of variation observed in our sample of 24 species from eight families. However, one species of Acanthodactylus (Lacertidae) had a consistent region of large tonic fibers (that did not stain darkly for either succinic dehydrogenase or myosin ATPase activity), a fiber-type only occasionally seen in the other 23 species examined. Many species have a large proportion of either fast-twitch glycolytic (FG; e.g. sand lizards and Aspidoscelis) or fast-twitch oxidative-glycolytic (FOG) fibers (e.g. horned lizards), with the slow-oxidative proportion occupying only 1-17% of the iliofibularis. Importantly, the negative relationship between FG and FOG composition observed in Phrynosomatidae appears to be a characteristic of lizards in general, and could lead to functional trade-offs in aspects of locomotor performance, as has previously been reported for Lacertidae. Reconstruction of ancestral trait values by use of phylogenetically based statistical methods indicates especially large changes in fiber-type composition during the evolution of horned lizards.

Animals↗

Analysis of monoamines, adenosine and GABA in tissues of the land snail Helix lucorum and lizard Agama stellio stellio during hibernation.

The aim of the present study was to determine the levels of monoamines, GABA and adenosine in the brain, heart and haemolymph of the land snail Helix lucorum and in the brain, heart and blood of lizard Agama stellio stellio during long-term hibernation. We measured levels of the monoamines serotonin (5-HT) and its main metabolite 5-hydroxyindole-3-acetic acid (5-HIAA), dopamine (DA) and its metabolites dihydroxyphenylacetic acid (DOPAC) and homovanilic acid (HVA), norepinephrine (NE) and epinephrine (E). The most abundant amines detected in the brain and heart of active H. lucorum were 5-HT and DA. Of the metabolites examined only 5-HIAA was found in the brain. NE was found at very low levels but only in the brain, while E was not detected in the brain and heart. The levels of 5-HT and 5-HIAA increased in the brain and heart of H. lucorum within the first months of hibernation, showing a significant decrease thereafter. The levels of DA did not change during hibernation. The results indicated that 5-HT might be involved in preparing snails for entry into hibernation. GABA was only found in the brain of H. lucorum, and the levels were low; these levels remained during hibernation. Adenosine was present in brain and heart of H. lucorum, and during hibernation, the level of adenosine decreased significantly in the brain but remained steady in the heart. The monoamines 5-HT, DA and NE were present in the brain of active lizards A. stellio stellio, whereas E was found only at very low levels. Moreover, the metabolites 5-HIAA, DOPAC and HVA were detected in the brain of active lizards. The monoamines 5-HT, DA, NE and E were also detected in the heart and blood of active lizards. During hibernation the levels of these four monoamines were decreased significantly in the brain and heart of A. stellio stellio. In contrast, the levels of E increased in the heart and blood of hibernating lizards. Adenosine was detected in both heart and brain of active lizards, but hibernation caused a marked decrease in its levels at both tissues. GABA was found at higher levels than monoamines and adenosine in the brain of active lizards, and hibernation caused a significant increase in its levels, indicating an important role of GABA in inhibition of neuronal activity in hibernating lizards.

Adenosine↗

Reservoir role of lizard Psammodromus algirus in transmission cycle of Borrelia burgdorferi sensu lato (Spirochaetaceae) in Tunisia.

To investigate the reservoir role of the lizard Psammodromus algirus for the Lyme disease spirochete, 199 lizards were trapped from April to October 2003 in El Jouza, northwestern Tunisia. In this site, the infection rate of free-living Ixodes ricinus (L.) by Borrelia was evaluated by immunofluorescence as 34.6% for adult ticks and 12.5% for nymphs. Eighty percent of P. algirus (117/146) captured during this study were infested by I. ricinus, the predominant tick species collected from lizards. The intensity of tick infestation of this host by larvae and nymphs ranged from 0.14 to 7.07 and from 1.5 to 6.58, respectively. These immature stages of I. ricinus were found on lizards in spring and the beginning of summer, with a peak of intensity during June (10.16 immature ticks by lizard). Tissue cultures from lizards and xenodiagnosis with larval I. ricinus were used to assess the infection and the ability, respectively, of infected lizards to transmit Borrelia to naive ticks. Seventeen percent of xenodiagnostic ticks (40/229) acquired B. lusitaniae while feeding on P. algirus. Therefore, we demonstrated the ability of the lizards to sustain Borrelia infection and to infect attached ticks, and we proved that P. algirus is a reservoir host competent to transmit B. lusitaniae.

Animals↗

Morphological evolution of the lizard skull: a geometric morphometrics survey.

Patterns of diversity among lizard skulls were studied from a morphological, phylogenetic, and functional perspective. A sample of 1,030 lizard skulls from 441 species in 17 families was used to create a lizard skull morphospace. This morphospace was combined with a phylogeny of lizard families to summarize general trends in the evolution of the lizard skull. A basal morphological split between the Iguania and Scleroglossa was observed. Iguanians are characterized by a short, high skull, with large areas of attachment for the external adductor musculature, relative to their sister group. The families of the Iguania appear to possess more intrafamilial morphological diversity than families of the Scleroglossa, but rarefaction of the data reveals this to be an artifact caused by the greater number of species represented in Iguanian families. Iguanian families also appear more dissimilar to one another than families of the Scleroglossa. Permutation tests indicate that this pattern is real and not due to the smaller number of families in the Iguanidae. Parallel and convergent evolution is observed among lizards with similar diets: ant and termite specialists, carnivores, and herbivores. However, these patterns are superimposed over the more general phylogenetic pattern of lizard skull diversity. This study has three central conclusions. Different clades of lizards show different patterns of disparity and divergence in patterns of morphospace occupation. Phylogeny imposes a primary signal upon which a secondary ecological signal is imprinted. Evolutionary patterns in skull metrics, taken with functional landmarks, allow testing of trends and the development of new hypotheses concerning both shape and biomechanics.

Animals↗

Evidence for orientation using the e-vector direction of polarised light in the sleepy lizard tiliqua rugosa

Adult sleepy lizards (Tiliqua rugosa) were trained to orient in a predictable direction under natural sky light in outdoor pens. When tested under clear skies in the late afternoon, without a view of the sun, the lizards exhibited a symmetrical bimodal pattern of orientation with respect to the trained axis. Since the e-vector of polarised light provides an axial rather than a polar cue, the bimodal orientation exhibited by the lizards is consistent with the use of a celestial compass based on sky polarisation patterns. To confirm that the lizards could orient with respect to a polarisation pattern, lizards were trained in indoor pens to orient in a predictable direction under a linearly polarised light source. When tested in a circular arena illuminated by another polarised light source, the lizards used the e-vector direction of the polarised light source to orient along the trained axis. There was no evidence that the lizards were using any room-specific cues or brightness patterns to orient in the training direction. These results support the hypothesis that the lizards can use the e-vector direction of polarised light in the form of a sky polarisation compass.

Journal Article↗

Interleukin-1 beta reduces daily activity level in male lizards, Sceloporus occidentalis.

In mammals, interleukin-1 (IL-1) mediates many of the behavioral consequences of pathogen infection. Other vertebrates show behavioral changes when infected, but the neuroendocrine bases of these changes are seldom known. Here we report that IL-1 beta alters the daily activity cycle of lizards (Sceloporus occidentalis) similar to that seen in lizards infected with malaria. To our knowledge, this is the first report of behavioral effects of interleukin in lower vertebrates. Male lizards were injected with human IL-1 beta (10 ng/g animal), saline, or nothing (control) and the activity level (proportion of lizards above ground) was monitored for 48 h. IL-1 beta-injected lizards showed a decrease in activity compared to saline-injected and control lizards within 2 h after treatment. Activity levels were equivalent among treatment groups during the middle of the day (1200-1600 h), but IL-1-treated animals were significantly less active in the morning (0800-1200 h) and evening (1600-2200 h). This decrease in activity is similar that seen in free-ranging lizards infected with malaria, supporting the hypothesis that IL-1 mediates the pathogen-induced alterations to activity behavior under natural conditions.

Animals↗

The Coachella valley fringe-toed lizard (Uma inornata): genetic diversity and phylogenetic relationships of an endangered species.

A phylogeny was reconstructed for 23 populations of fringe-toed lizards (genus Uma) from the three most northern species of the genus, including the Mojave fringe-toed lizard U. scoparia, the Colorado Desert fringe-toed lizard U. notata, and the endangered Coachella Valley fringe-toed lizard U. inornata. The outgroup taxa were the zebra-tailed lizard, Callisaurus draconoides; the lesser earless lizard, Holbrookia maculata; and the greater earless lizard, Cophosaurus texanus. Evaluation of 1630 combined nucleotide sequence from the mitochondrial genes ATPase 6 and cytochrome b yielded 10 most parsimonious trees. Reweighting the characters using the rescaled consistency index eliminated eight of these trees. The remaining two trees differ only in the placement of two individuals from the Superstition Mountains which either formed a monophlyetic unit or grouped with one individual from the Anza-Borrego population. The preferred phylogeny, one more consistent with geography, had two primary clades: one consisting of U. scoparia and the other placing U. inornata inside the clade containing U. notata. Uma inornata was most closely related to nearby U. notata notata, as opposed to more distant U. notata rufopunctata.

Adenosine Triphosphatases↗

A juvenile lizard specimen with well-preserved skin impressions from the Upper Jurassic/Lower Cretaceous of Daohugou, Inner Mongolia, China.

Lizards are now relatively well known from the Yixian Formation of northeastern China. In this study, we describe a juvenile lizard from a fossil horizon at Daohugou, Inner Mongolia. These beds predate the Yixian Formation, and are probably Late Jurassic or earliest Cretaceous in age. The new specimen thus documents the first lizard material from the Daohugou locality and is the earliest lizard skeleton from China. Comparisons with developmental stages of modern lizards suggest the Daohugou lizard is a hatchling. Although tiny, the specimen is notable in preserving exquisite skin impressions showing the variation in scalation across the body, the shape and position of the cloacal outlet, and details of the manus and pes. These are the earliest recorded lepidosaurian skin traces. In its general proportions and the possession of paired frontals, the small Daohugou lizard resembles both the Yixian taxon Yabeinosaurus tenuis and the questionable Jeholacerta formosa, but it differs from the latter in scalation and, based on other characters, may be distinct from both.

Animals↗