A morphology-based taxonomic revision of Laudakia G ray , 1845 (Squamata: Agamidae)

The former genus Stellio has already been partitioned into Laudakia G ray , 1845 and Acanthocercus F itzinGer , 1849 on the basis of several pieces of evidence. The main objective of this study is to revise Laudakia which recently includes 20 species: L. agrorensis, L. badakshana, L. bochariensis, L. caucasia, L. dayana, L. erythrogaster, L. fusca , L. himalayana, L. lehman­ ni, L. melanura, L. microlepis, L. nupta, L. nuristanica, L. pakistanica, L. papenfussi, L. sacra, L. stellio , L. stoliczkana, L. tuberculata, and L. wui . More than 600 specimens have been studied with reference to 54 morphological characters which resulted in a detailed descriptive account for each taxon. Agama isozona is recognized as a synonym of L. bochariensis. The latter species itself has been placed in a supraspecific complex consisting of L. himalayana, L. badakshana and L. bo­ chariensis. Laudakia caucasia which has ben lowered and raised several times since its appearance is again identified as a monotypic species by placing L. caucasia triannulata as synonym under L. microlepis. Laudakia fusca was described as a variety of L. nupta but subsequent herpetologists synonymized it or recognized it as full species. According to this study L. fusca should be recognized again as subspecies of L. nupta pending more detailed further research. Moreover, several previous works have indicated that Laudakia is paraphyletic and therefore two new genera are described herein encompassing the stellio - and caucasia -groups.


Prologue
The present paper is based on the PhD thesis of the late Dr. Khalid Javed BaiG who passed away through a tragic accident on November 11 th , 2006 (see the obituary for him in the Russian Journal of Herpetology 14 [1], 2007).Khalid had been awarded with a fellowship by the German Academic Exchange Service (DAAD) from October 1990 to March 1992 at the Herpetology Section of the Zoologisches Forschungsmuseum A. Koenig in Bonn, Germany.During this time he prepared and finished his thesis [supervised by the fourth author (WB)].It was defended subsequently at the University of Islamabad, in his home country Pakistan.We think that Khalid J. BaiG's thesis is a very important contribution to the knowledge of agamid lizards, particularly of the whorl-tailed agamas of the genus Laudakia.Therefore, we decided to complete and update the manuscript after some years with the most important data and references published in the meantime, wishing to prevent this major contribution from suffering the same fate of another famous thesis on agamid lizards (Moody 1980) that unfortunately remained unpublished until today.The third author (NBA) was also cooperating with Khalid and still is a dedicated researcher on agamids.This is also true for the second author (PW) who organized the first international symposium on agamid lizards in Bonn, DeAgamis I, in 2008 which was followed in 2010 by DeAgamis II in St. Petersburg, organized by NBA.Both these meet-

Introduction General Introduction
Taxonomically, the history of Laudakia dates back to 1758 when the first member of this group was described as Lacerta stellio by linnaeus.At that time knowledge of systematics was not that advanced and the morphological and behavioral similarities of agamids and iguanids misled many herpetologists to confuse their affiliations.As example, the forementioned species was also described as Iguana cordylina by laurenti in 1768.These systematic anomalies were even greater concerning iguanids.The present Phymaturus palluma (Molina, 1782) was identified by its author as Lacerta in 1782 but placed in Stellio in 1801.Microlophus peruvianus (lesson, 1830) was firstly described as Stellio in 1830 whereas Uracentron azureus was first described as Lacerta azurea by linnaeus (1758) and was placed in Stellio in 1801.Likewise, Tropidurus torquatus was described in 1820 as Stellio torquatus (Burt & Burt 1931).
BoulenGer (1885) resolved many discrepancies at least at family level and fixed several taxa into their respective families and genera.Among the Agamidae, which is the fourth largest family among lacertilians and distributed from Africa through Asia to Australia, Stellio has long been a controversial group.Many herpetologists considered it as part of the African genus Agama daudin, 1802(e.g. BoulenGer 1885;sMith 1935;WerMuth 1967).Although the name Stellio laurenti, 1768 was made unavailable by steJneGer's (1936; see also henle 1995) action of selecting the non-identifiable Stellio saxatilis as the type species, several herpetologists kept it alive.More recently Moody (1980) resurrected six distinct genera from the collective genus Agama, including Stellio.Moody (1980) placed 22 species into the genus, including several African taxa.Since then, successive herpetologists have largely accepted the importance and individual status of this group.Some of them recognized it as an independent genus Stellio (ananJeva & ataev 1984;ananJeva & danov 1990;ananJeva et al. 1990ananJeva et al. , 1991;;osneGG 1989;JoGer & arano 1987) while others treated it as an informal group within Agama (BaiG & BöhMe 1991a, b;BöhMe 1981;JoGer 1991).BaiG (1992) and BaiG & BöhMe (1997) partitioned Stellio (sensu Moody 1980) into the genera Laudakia Gray, 1845 and Acanthocercus FitzinGer, 1849.Their morphological analysis was based on 54 characters and incorporated anatomical, karyotypic and biochemical evidence from relevant literature.
A genetic study of the entire genus is lacking and is now under the focus of the authors.However, some groups of Laudakia are comparatively well studied, e.g. the Laudakia caucasia species group (MaCey et al. 1998(MaCey et al. , 2000b) ) and species of Central Asia (Melville et al. 2009), or laudakias were used to solve zoogeographical questions (MaCey et al. 2000a).
However, one of these studies (MaCey et al. 2000a) has shown that Laudakia, as recently recognized, is paraphyletic.They recognized the L. caucasia group as sister taxon to a clade containing some Phrynocephalus species and Laudakia stellio, whereas the L. tubercu lata group is basal to this entire clade (see fig. 1).Even if these bootstrap values (see fig. 1) are not all highly significant, the results are supported by WaGner (2010) who also recognized Laudakia as paraphyl-2009) the same groups as mentioned above are in distinct clades, which is supported by some morphological characters (BaiG 1992).Very recently, edWards & Melville (2011) published a study including mtDNA analyses showing Laudakia as monophyletic and sister taxon to Phrynocephalus (fig.2).Also here L. stellio is isolated and sister to the L. caucasia group, while the L. tuberculata group is basal to the previous groups.In all these different studies the same groups are supported, but there is evidence of a paraphyly of the genus.Therefore, we decided to classify the different species groups in distinct genera.

Previous Works
Karyotypic studies on agamids have been carried out by hall (1970), GorMan & shoChat (1972), GorMan (1973), soKolovsKi (1974), KuPriyanova (1984).In addition, Moody & hutterer (1978) and Witten (1978) described karyotypic formulas for several agamids.These studies suggest that Laudakia possess 2n = 36, a pattern considered ancestral for all lizards on the basis of its occurrence in nearly all lizard fami-etic in a study additionally including Acanthocercus, Xenagama and Bufoniceps.In contrast, Melville et al. (2009) showed Laudakia as monophyletic with the exception of Xenagama batillifera which was part of the laudakian clade, which should be a result of a misidentification as the other Xenagama species is a sister taxon of Agama in the same study.Moreover, the monophyly of Xenagama is supported by WaGner (2010).However, also in the previous study (Melville et al.  sieBenroCK (1895), duda (1965) and ananJeva (1980) categorized the teeth as incisors, canines and molars, whereas Moody (1980) referred to the anterior pleurodont teeth as canines and the posterior acrodont teeth as molars.The first three authors invariably reported one incisor and one canine in each quadrant of the maxilla and dentary in all species of the genus Agama at this time (sensu WerMuth 1967).
Behavioural patterns of agamids are very similar to those of iguanids and until we have more information on the social behaviour of additional species of agamids, the possibility of significant differences between the social behavioral patterns of these two families remains an open area of investigation (BlanC & CarPenter 1969).BrattstroM (1971) described some 73 behavioral postures and positions for Australian bearded dragon, based on observations made in the field and the laboratory.Information regarding agamid spacing systems is sparse, but the few studies carried out suggest that agamids are remarkably similar in habits and social structure to iguanids (staMP 1977).sMith (1935( ), harris (1964( ), sChMidt (1966)), BlanC & CarPenter (1969), BrattstroM (1971), CarPenter (1978), sChleiCh (1979), orlova (1981a, b), Beutler (1981Beutler ( ), daniel (1983) ) and many others have suggested territorial behavior in agamids.These authors mostly attribute territoriality to the males.Only sChMidt & inGer (1957), Madel & KloCKenhoFF (1972) and lanGerWerF (in orlova 1981a) observed it in female agamids as well.Moreover, Panov & zuKova (1995) studied the variability and differentiation of e.g. home ranges in populations of L. caucasia.

Material
Pakistan Museum of Natural History, Islamabad and Zoologisches Forschungsmuseum Alexander Koenig, lies (WilliaM & hall 1976), a finding which is also observed in Uromastyx, Leiolepis, Physignathus and Acanthocercus.
Cranial morphology and dentition was studied by CaMP (1923) who reviewed a number of useful morphological characters that vary at the subfamilial and generic levels, but he examined few agamids.Jollie (1960) succinctly reviewed the head skeletons of several lizards, including agamids.However, both authors relied largely on German-speaking morphologists, primarily sieBenroCK (1895) who had depicted skulls and mandibles of several agamid species including L. himalayana and L. tuberculata.Moody (1980), using 78 characters, emphasized the skeletal characters in building phylogenetic relationships among genera of the Agamidae.ananJeva (1980) published structural characteristics of the skull, dentition and hyoid of L. caucasia, L. erythrogaster, L. himalayana, L. lehmanni, T. ruderatus and Trapelus sanguinolen tus.Besides el-touBi (1947) and duda (1966) who studied L. stellio and L. tuberculata respectively, other information about this group is scattered and mostly concerned with non-Stellio (sensu Moody 1980) members of Agamidae (sMith 1935;Barry 1953;GeorGe 1955;harris 1963).Also ananJeva (1980) studied skull and associated structures of six agamids, with four belonging to the stellio-group.
The agamids are conventionally characterized as the lizards with acrodont teeth (roMer 1956).This characterization is too general and partly false (see Moody 1980;sMirina & ananJeva, 2007).Agamid teeth are heterodont i.e., comprise more than one form of teeth, viz.anteriorly pleurodont and posteriorly acrodont.Description of agamid dentition has also been given by several authors (edMund 1969, CooPer et al. 1970, PresCh 1974, roBinson 1976) but Moody's (1980) point of view seems to be more reasonable.Moody (1980) explains the differential tooth size from anterior to posterior by stating that anterior acrodont teeth are ankylosed juvenile teeth which could not increase in size.The pleurodont teeth at the tip represent an intermediate evolutionary step to higher forms because they do not undergo replacement at regular intervals (except in case of accidental loss) and have partial basal ossification.sieBenroCK (1895) reported two pleurodont teeth in Agama hispida, Laudakia himalayana, Laudakia stellio, Trapelus mutabilis pallidus and Trapelus sanguinolenta, and three in Laudakia tuberculata, but only one in Agama atra and Agama agama. ananJeva (1980) mentioned two in Trapelus ruderatus (now recognized as T. lessonae fide rasteGar-Pouyani 2000) and variably two or three in Trapelus sanguinolenta, L. caucasia, L. lehmanni and L. erythrogaster. duda (1965) and BaiG (1991) confirm the tooth count given by sieBenroCK (1895) for L. tuberculata.
Vertebrate Zoology n 62 (2) 2012 tion, and important skulls are presented as drawings, which were made with the help of a camera lucida.

General Morphology
There is no significant difference between the number of supralabials and infralabials in all members of Laudakia.In some taxa, e.g.L. stoliczkana, there are slightly more supralabials, whereas L. stellio vulgaris shows more infralabials.Most species have nine to twelve supralabials.However, L. nupta and L. eryth rogaster have higher numbers.The lowest number of supralabials is represented in L. sacra and L. agroren sis some of whom have below nine.
In general the number of lamellae under the third finger is smaller than under the fourth toe.In most taxa this difference is approximately five scales, but in L. dayana, L. stoliczkana, L. erythrogaster, L. hima layana and L. nuristanica, it is higher.The highest numbers can be found in L. dayana and L. nuristani ca.Under the third finger the number usually remains smaller than 20 and under the fourth toe smaller than 25.Specimens of L. tuberculata, L. nuristanica and L. dayana may exceed 30 and sometimes reach even 35 subdigital lamellae.
In some cases the number of scale rows around midbody is an important character to distinguish different of the herein mentioned taxa.Laudakian species mainly stay within the range of 100 -150 scale rows around midbody.However, the minimum numbers (i.e. less than 100) are recognized in L. nupta, L. erythrogaster and L. lehmanni, whereas the highest numbers in L. nuristanica.This latter species shows more than 200 scales around the body.Among others especially L. tuberculata, L. microlepis and L. s. picea also show a fairly high number of scales rows around midbody.
In some cases the number of pericaudal scales reflects a similar pattern as the number of scales around the body.L. nuristanica, L. tuberculata, L. stoliczka na, who all have a high number of scales around midbody, also show a high number of pericaudal scales, while in L. s. picea has low numbers in both characters.The largest caudal scales can be found in L. mela nura, but also Laudakia s. cypriaca and L. nupta show relatively large scales.
In body proportions, the longest tails can be found in L. dayana and L. melanura, whereas the short-Bonn are the primary sources of information for the present study, but in addition almost all other major Museums of Europe and one in the USA have been visited to examine the material deposited there.The list of these institutes with their acronyms (used to quote these institutions in the following text) is as follows: We have studied almost all related material housed in these institutes.The selected number was only studied when there were several specimens from the same locality.The data have been compiled based on 54 morphological characteristics found important for the comparison of the different members of this group (see BaiG 1992).Of these, 22 are morphometric, 7 quantitative numeric and 25 qualitative (see BaiG 1992).Measurements were taken nearest to the 10th of a millimeter with the help of a vernier calliper.The scales on the different parts of body are counted using Stereomicroscope.All statistical data has been analysed on the computer using Microsoft Excel (see BaiG 1992).width.The other taxa in which tail width exceeds one and half times the tail height are L. agrorensis, L. bo chariensis, L. caucasia, L. melanura, L. microlepis, L. pakistanica, L. s. stellio and L. tuberculata.The least depressed tail have been found in L. s. cypriaca, L. lehmanni and L. nupta.The proportional length of the upper arm (humerus) is always higher than the lower arm (radius and ulna).With reference to the hand, the humerus is only slightly longer in L. mela nura, L. nupta and L. stellio picea and almost equal in L. s. brachydactyla and L. erythrogaster.In all other species, the hand is longer than the humerus.The hand is always longer than the lower arm, although L. mela nura and L. nupta show a relatively longer lower arm compared with other Laudakia species.The ratio of hand length to the length of the third finger and the ration of foot length to the length of the fourth toe do not show any significant variation among all the taxa.In the Hindlimb the thigh is always longer than the shank, but with reference to hand it is equal or more in L. melanura, L. nupta, L. s. picea and L. s. stellio.In all other taxa, the foot is longer than the thigh.The shank is always shorter than the foot, but again L. me lanura and L. nupta show relatively longer shanks as compared to other members of Laudakia.est in L. erythrogaster, L. microlepis and L. s. picea.Lau dakia stoliczkana has the shortest head, whereas the longest heads can be found in all subspecies of L. stellio.The other species show more or less the same range regarding the head length.Laudakia tu berculata, L. agrorensis and L. pakistanica show the least relative head height.The width of head is more or less one and half times that of its height in most taxa, although L. agrorensis shows the highest proportion.The ratio of the snout length to the distance between eye-tympanum indicates that eyes are always located more towards tympanum.In L. agrorensis, L. dayana, L. nuristanica and L. tuberculata the eye is closer to the tympanum than in the other members of Laudakia.The ratio eye-width to tympanum diameter is unique in L. melanura and L. nupta, where eyewidth is smaller than tympanum diameter.In all other taxa the ratio is reversed.In case of limbs, although the tendency is more or less similar to head length, L. stoliczkana altaica, L. lehmanni and L. sacra show relatively shorter limbs than other taxa.The ratio of forelimb to hindlimb is almost uniform in all members of Laudakia.In all species, the tail width exceeds tail height.Laudakia s. picea has the most depressed tail, where the height of the tail is about half of its articulation with surrounding elements and provides principal support for the lacrimal duct.The base of the lacrimal is more robust, not visible externally and firmly sutured between the palatine and maxilla in the anterior orbital floor.

Cranial morphology and dentition
The postorbital is large, forms much of the posterior margin of the orbit and lies in a horizontal plane with a slight ventrolateral tilt.It forms the temporal arch and, together with the squamosal, the lateral margins of the supratemporal fossa.The dorsoanterior margin of the postorbital defines the posterior apex of the orbit.Here, it articulates with the lateral processes of the frontal and parietal.The jugal forms the entire ventral margin of the orbit without any participation of the maxilla.It has a strong dorsal process that forms much of the anteroventral part of the temporal arch and acutely inserts between the postorbital and squamosal.The third pair of skull opening, the temporal openings, are each surrounded by the postorbital that forms much of the posterior margin of the orbit, parietal, supratemporal and squamosal bones.A single parietal bone covers most of the upper skull.The pineal foramen is near the anterior border of the parietal bone whereas at its posterior border two long supratemporal/parietal processes pass back diagonally towards the squamosal and quadrate.The origins of the supratemporal processes are widely separated and project posteriorly, curving only slightly below the horizontal plane.A downwards projection of the anterior base of the supratemporal process of the parietal produces a partial lateral wall to the cranial case.
The supratemporal is a small splint that closely adheres to the ventrolateral edge of the parietal process and articulates by means of a triangular-shaped head with the dorsoanterior surface of the quadrate, paroccipital process and the squamosal.It completely separates the last two elements from mutual contact.The squamosal is the posterior element of the temporal arch.It forms an overlapping joint with the postorbital and jugal anteriorly and a hinge joint with the dorsal head of the quadrate posteriorly.The foramen magnum, through which the spinal cord passes, is surrounded by four bones: dorsally by the supraoccipital, ventrally by the basioccipital and laterally by two exoccipital bones.The supraoccipital is a broad, hour-glass shaped and fused midline element which continuously contacts the posterior cranial wall of the parietal.The basioccipital forms the posterior floor of the brain-case and contributes to the basal portion of the occipital condyle.
The exoccipital bones are paired elements lying on either side of the foramen magnum contributing to the occipital condyle.The palatal region comprising denticulate and non-denticulate elements and is observable from the ventral view of the skull.Anteriorly, there are two crescent shaped openings, the internal nares,

Cranial Morphology
Variation among different bony elements are generally noticeable and significant at the generic or suprageneric level.Specific level is typically too low to contribute something significant in this regard.However, to demonstrate different elements and features of the skull and associated structures, drawings of some specimens are being presented (figs. 3a, b).The skull of Laudakia is depressed and triangular when viewed from above.From the dorsal side three pairs of openings or vacuities are visible, i.e. nasal openings, orbital openings and temporal openings.The nasal openings are small and lead through the olfactory chamber to the buccal cavity.They are separated from each other by the premaxilla and bordered laterally by the maxilla and posteriorly by the nasal bones.The premaxilla is a single midline bone bearing only pleurodont teeth.It is T-shaped with an arching denticulate margin and a long internarial shaft that forms an arching buttress of the snout and a firm overlapping joint with the compressed anterior process of the nasals.The maxilla is the major tooth-bearing element of the cranium.The anteriormost two teeth are enlarged and pleurodont.The remainders are acrodont.The maxilla continues posteriorly to the last tooth as a sharp process, dorsally sutured with the jugal and medially overlapped by the ectopterygoid.The palatal portion of the maxilla comprises a narrow shelf that expands anteriorly to form a suture at the midline and overlap the palatal portion of the premaxilla.The nasals are invariably paired and in contact along the midline.Posteriorly, they overlap the frontal.Laterally, as a thin process, they overlap the dorsal process of the maxilla and border the prefrontal.The two orbital openings are separated by a median frontal bone.Their anterior border is formed by the prefrontal bone, a small lacrimal bone and the maxilla, whereas their posterior border by the postorbital and jugal bones.The frontal is posteriorly broad, due to the lateral processes that extend to the postorbitals.The suture with the parietal is thus broad and straight and bends slightly posterior at the midline where it forms the anterior margin of the parietal/pineal foramen.Anteriorly, it is doubly fork-shaped.A midline process separates the nasals posteriorly and contacts the internarial process of the premaxilla beneath the nasal and lateral processes.On both sides it separates the prefrontal and nasal.The prefrontal bones are robust and form much of the anterior rim and wall of the orbit.The dorsolateral corner of the prefrontal is the apex of the dorsal and lateral surfaces of the snout, and it is also the posterior culmination of the rostral canthal ridge.The lacrimal forms the anteroventral margin of the orbit and a small portion of the posterolateral snout region.It is a thin plate without firm functional leading from the nasal cavity, separated by the vomers and bounded at their hind end by the palatine.The palate itself is bounded at its anterior tip by the single premaxilla and along its lateral border by the maxilla.These two bones constitute the denticulate elements of the palate and have already been described under the heading of nasal openings.Vomer, palatine, pterygoids and ectopterygoids are said to be non-denticulate elements of the palate.Vomers are thin anteriormost elements of the palate and lie between the internal nares.The latter are paired and their medial and lateral borders turn dorsally.The palatines are the middle elements of the palate and form most of the medial margin of the fenestra.The posterior part of the internal narial margin is formed by the palatine.The suture with the vomer is transverse and lies more closely to the posterior margin of the internal nares.The palatines are paired and form a loose ligamentous contact along the midline.The palate is not a flat roof of the mouth but instead bends midway along the palatine to form a horizontal anterior margin.The floor of the nasal capsule comprises the vomers and anterior palatines.The pterygoids constitute the largest element of the palate.The posterior process connects with the quadrate and the braincase via the basisphenoid.The anterior palatal portion is firmly sutured with the palate and adjacent cranium.A ventrally projecting process has sliding contact with the coronoid process of the mandible.The pterygoid do not suture on the midline, however, a membranous contact across the interpterygoid space is very likely.The medial margin of the pterygoid, which lies on the floor of the palate, is divergent posteriorly.The suture between the pterygoid and palatine is firm.The pterygoid forms a strong downward projecting process, which articulates in a sliding manner with the coronoid process of the mandible.The ectopterygoids bridge the pterygoid, the posterior part of the maxilla and the anterior part of the temporal arch.Along the internal margin of the posterior wall of the orbit, a dorsal process of the ectopterygoid contacts a ventral process of the postorbital.The medial process of the ectopterygoid overlaps the pterygoid dorsally.The ectopterygoid forms the posterior margin of the palatine fenestra and also part of the lateral margin due to the anterior process running medially to the infraorbital process contributed by the jugal.

Mandible
Seven bony elements, namely the dentary, angular, supraangular, articular, prearticular, splenial and coronoid, constitute the mandible and may also be termed as mandibular elements.The dentary is the largest of these bones and extends labially to the posterior coronoid process and almost reaches the articular.
Lingually it is larger than the small splenial and the splint-like angular which curves along the ventral margin of the Meckelian groove to the ventral edge.The dental gutter in which the dental papillae are situated does not extend very deeply along the lingual surface of the dentary and is nearly straight.Posteriorly, the dentary strongly overlaps the supraangular as a sharply pointed dorsal and ventral process.Within the acute notch formed by these processes lies the anterior supraangular foramen.In all examined material the angular is always large and contains an angular foramen medially.The foramen is located below the anteromedial coronoid process.The supraangular is visible labially where it is overlapped by the dentary and forms an acute margin and suture with the angular.An anterior foramen is present in the apex of the notch produced by the dentary process.Lingually, the supraangular can be seen forming the internal wall of the mandibular fossa.The articular is the only endochondral bone of the mandible.It is indistinctly fused with the dermocranial prearticular bone.It forms the articulating facet for the quadrate and the retroarticular process on which the depressor mandibular musculature inserts.The prearticular is on the lingual surface of the mandible and extends from the articulating surface of the articular to the anteromedial process of the coronoid.It forms the lingual or internal wall of the mandibular fossa.Ventrally, it forms an acute border with a distinct suture to the angular.The splenial is present as small bone flakes.It is flat, thin and roofs the posterior part of the Meckelian canal.It overlaps the dentary, the anteriomedial process of the coronoid, the prearticular and the angular.The large coronoid has a prominent dorsal process which articulates as a sliding joint to the opposing coronoid process, formed by the pterygoid and ectopterygoid.The wide coronoid dorsal process curves slightly posteriorly.The lingual surface of the coronoid is an inverted V with well defined anterior and posterior medial processes.The anterior process is flat and overlapped by the dentary and the splenial.The posterior process is ridged and compressed in a transverse plane near the tip which overlaps the prearticular.

Dentition
Two types of teeth can be found amoung species studied in this publication.The acrodont tooth has a broad and slightly swollen basal portion which is ankylosed against the medial wall and floor of the shallow dental gutter.The shearing portion is triangular and compressed laterally, with the labial surface more flattend than the lingual surface.The pleurodont teeth are large and conical.The premaxilla has three teeth in L. agrorensis and L. pakistanica and apparently two in L. himalayana.The maxilla carries anterior two pleurodont teeth behind which the number of acrodont teeth ranges from 12 -14.The dentary shows a similar pattern of pleurodont and acrodont teeth.The number of pleurodont teeth varies between two and three.There are two in L. pakistanica and L. himalayana and three in L. agrorensis.The number of acrodont teeth (molars) ranges between 14 -15.

Species Accounts
The systematic account of the species, based on all 54 morphological characteristics analysed by BaiG (1992), has been given separately for each species.It begins with the synonymy of each taxon, followed by diagnosis, description and distribution.Important aspects of previous studies, together with own observations and comments, have been included in `Remarks' at the end of each taxon chapter.Murthy (2010) published a book about the reptiles of India, classifying Brachysaura minor as Laudakia.However, he only mentioned the species in a checklist and a short species account, but he failed to explain his re-classification.Therefore, we still recognize this species as Brachysaura and further research will show the correct position of this taxon.
In general, all herein mentioned lizards can be characterized as diurnal, conspicuously active occupants of terrestrial mountainous habitat (with the exception of L. stellio), and are visually oriented in feeding and social behavior.In morphology, physiology, and behavior they show several characteristics which may be observed in other agamids and iguanid taxa.Laudakia lizards possess a head and body more or less depressed; tympanum distinct, diameter half or more than that of eye; groups or series of spinose scales on neck and around tympanum; nuchal crest absent or represented by a row of spinose scales; gular sac absent (some species show slight tendency); gular plicate; head scales heterogeneous; vertebral scales usually enlarged; femoral glands absent; callous glands present in males (in some species also in females); tail oval in cross section; and caudal scales form distinct annuli.As typical agamid lizards they have strong limbs which aid efficient in running on the ground and climbing on the rocks and long, slender, oval, tapering tails which is not capable of autotomy like in lacertid lizards.However, it may be regenerated in case of accidental loss, regenerated mostly clubshaped, but sometimes bifurcated (BaiG 1988b;ananJeva & danov 1990) or elongated (WaGner et al. 2009).They may use the tail for defense during fighting with one another.Most members of this group are semiherbiv-orous, feeding on insects and plants (Moody 1980) Stellagama s. picea is the smallest among all subspecies and it is unique in its black colour with yellow dots.S. s. vulgaris is a medium sized lizard of the group and may be differentiated from other members by being dull brown in colour with broken vertebral patches and having a segmented tail with two whorls anteriorly and three posteriorly on the dorsal side, while two whorls occur ventrally.The complete change from two to three whorls per segment is characteristic for S. s. cypriaca.daan (1967) mentioned that S. s. brachydactyla was the largest but observations made by oseneGG (1989) and BaiG (1992) prove S. s. cypriaca as the largest subspecies.Stellagama s. brachydactyla is unique in its wide vertebral zone of similarly sized scales and is usually bright in colouration, sometimes with a reddish tinge.
Despite the efforts of many herpetologists, the taxonomic status of different populations of S. stel lio is still unclear.FloWer (1933) was the first who identified three different populations in the Egyptian-Palestinian region.haas (1951a) mentioned striking geographical differences and later he described (haas 1951b) a new subspecies of S. stellio from the Negev (southern Israel).ParKer (1934ParKer ( ), daan (1967) ) and Beutler & Frör (1980) subsequently described new races of S. stellio whereas BaiG (1992) recognised only four subspecies in addition to the nominate form.
Remarks.The groups of the whorl tailed agamas are distributed mainly an Asia while S. stellio is the only species occurring in southern Europe, northern Africa and on several different islands.All subspecies are pri-marily rock dwelling (some go on trees and buildings) at quite low elevations.
Stellagama stellio brachydactyla (Haas, 1951)  Stellagama stellio cypriaca (Daan, 1967) 20 -26 (23 ± 2.0) scales in first complete whorl around the tail.Callous glands present in males at precloacal and also at abdominal position, number of rows at precloacal position 3 -5.A very narrow, 2 -3 scales wide patch on abdomen.Head ash grey; gular also grey, rarely with few black scales or faded pattern; belly pale yellow but may be speckled; yellow vertebral blotches hardly visible in grown specimens but may be visible in juveniles, generally space between transverse enlarged scale rows filled with black, otherwise grey; enlarged transverse vertebral scales and other groups of enlarged scales light; tail light in proximal half and cross-banded distally.

Distribution. Endemic to Cyprus.
Habitat.According to Baier et al. (2009) a variety of dry habitats like stone walls, rocks, walls of old houses or on trees.
Stellagama stellio daani (BeutLer & Frör, 1980)  Colouration characterized as head coloured dark grey to black and not distinct in colouration from back; gular spotted or speckled with black, usually over half the area; belly whitish and pale speckled; 4 -5 yellow vertebral bloches on dark background, white spots on the flanks; tail whitish below, dark grey and and yellow bandes above.
Taxonomy.The status of Stellagama s. daani was unclear, because of doubts presented by BaiG (1992), but the study of alMoG et al. (2005) clearly shows significant differences in morphology between this subspecies and the nominate form.Also the different colouration of the head, in comparison to S. stellio, indicates at least a subspecific differentiation.
Distribution.Central Macadonia, central Cyclades, Saloniki, islands in the Aegian Sea and Turkey.
Studies by Beutler &Frör (1980) andalMoG et al. (2005) were uncertain concerning this range boundary.alMoG et al. (2005), Baran & öz (1985), Baran & atatür (1998) and GöçMen et al. (2003) accepted the occurrence of this subspecies in western Anatolia but assigned the population of south-eastern Anatolian coast to S. s. stellio.However, because they did not report the typical yellow to red head colouration of the nominate form and because alMoG et al. (2005) did not find significant differences between these populations and S. Stellagama stellio picea (Parker, 1935) 1935 Agama stellio picea ParKer, Proc.Zool.Soc., London, 1935: 137;pl Diagnosis.Broad band of enlarged vertebral scales, body scales heterogenous, tail segment of two scale rows.Differs from the similar S. s. brachydactyla as follows: band of somewhat enlarged vertebral scales with transverse rows of greatly enlarged scales, separated by smaller scales, enlarged scales on the left and right side; longer toes with an average of 18.5 subdigitalia; dorsal colouration with light markings forming numerous narrow transverse bars.This taxon differs from Stellagama s. vulgaris in having dorsal enlarged scales over the pelvis juxtaposed in transverse rows.
Description.Head moderately heavy, less depressed; snout longer than the distance eye-tympanum or eye width; tympanum exposed, more than half of eye width; nostril pierced below canthus rostralis, equal or less than half of the size of the nasal scale, touching rostral or sometimes interrupted by one scale; no true gular pouch but occasionally show some tendency, especially in males; gular strongly plicate; upper head scales heterogeneous, subequal, usually smooth above but may be rugose posteriorly; labials 9 -13 (11.6 ± 0.91); groups of spinose scales present on the neck and sides of head especially around tympanum; vertebral zone of enlarged scales, vertebral scales heterogeneous, irregular, usually smooth or weakly keeled, larger than other small dorsals, 16 -19 transverse rows of enlarged mucronate scales between axila and groin; other small dorsals distinctly smaller than enlarged ones; ventral scales smooth, smaller than enlarged vertebral; gular scales mucronate or spinose; limbs very strong, covered with enlarged mucronate scales, hind-limb slightly longer than the distance between gular fold and cloaca; fingers and toes compressed, 16 -23 lamellae under 4th toe.Tail moderately depressed, oval in cross section; distinct tail segment, each consists of 2 whorls of enlarged mucronate scales but may be mixed with a third in the terminal part of the tail; 3 -5 rows of callous glands with 25 -60 femoral pores present in males at precloacal.Ground-coloured grey, mid-dorsal enlarged scales orange, remaining enlarged scales metallic bluish grey, clusters of tubercles at the flanks cream-yellow, earhole and eye framed orange, hindlimbs and tail cross-banded yellow.Females lacking the bluish component of the enlarged dorsal scales.

Distribution. Southern Sinai and southernmost Israel.
Remarks.This recently described subspecies was recognized by biometric studies of the hardun in the Sinai and Negev deserts.The results indicated that the Sinai population differs from other populations and it was thus decribed as a new subspecies by Werner (in laChMann et al. 2006).This distinction coincides with biogeographic breaks in other taxa, such as Mesalina bahaeldini, Eirenis coronella ibrahimi and Hemidactylus mindiae.
Stellagama stellio stellio (Linnaeus, 1758) A wide variation of colour combinations, sometimes because physiological color change, is presently associated with the nominate form and future studies are required to reach any conclusions.However, the subspecies may be characterized, in preserved state, as head pale yellow to brownish grey; gular may or may not spotted or speckled with black; belly pale yellow but may be speckled; 4 -5 yellow vertebral bloches on dark background, enlarged transverse dorsal rows and other groups of enlarged scales usually yellow; tail light in proximal half and cross-banded distally.During breeding time or while defending terretories the males possess typical yellow to red heads.
Distribution.Greece, several islands of Cyclades, Turkey, Syria, Lebanon, Israel (excluding southern part) and western mountain regions in Jordan.
Habitat.ClarK et al. (1973), in their herpetological studies of Turkey, said that it is an inhabitant of rocky places, either natural or artificial (stone walls, bridge parapets, ancient buildings).However, they also observed them in a few unusual habitats: around holes in an earth ditch by the roadside, the base of bushes, and even more surprisingly between Gazipasa and Anamur several of them were seen up in olive trees in a field.Colour dull brownish grey, pattern of vertebral blotches is present in juveniles but in adults it mostly fades away and may be represented by some broken, irregular whitish spots; gular region dark grey with yellow ocelli which are more concentrated near tip or below the labials; belly yellowish brown and may be spotted.

Distribution. Northeast Egypt, in and around
Alexandria and Cairo, and probably coastal Sinai.
Remarks.This Levantine lizard is referred to as the Starred agama by FloWer (1933) and is identified as "Hardun" in Egypt.Among all 'Laudakia' in the old sense this is the first to be described by herpetologists.

Type species. Stellio caucasius eiChWald, 1831
Diagnosis.Tail arranged in distinct whorles of usually three, only sometimes two or four scale annulis.Gular scales smooth.Vertebral scales larger than other dorsal scales, body scales heterogeneous, irregular and keeled.Tail length about two times of the snout-vent length or longer.
Etymology.The chosen name refers to Laudakia (as allusion of the formerly used name of the genus) and para [greek for 'next to'] (to show the relation between the two genera).
Paralaudakia caucasia (eicHWaLD, 1831)  ClarK et al. (1969) identified some Afghan specimens with very high scale counts and showed their concern about S. microlepis.Even later, ClarK (1990) casted doubts about some specimens from north of Hindu Kush.BaiG (1992) also studied several specimens from Afghanistan and was convinced that both P. caucasia and P. microlepis exist there.However, it has yet to be determined whether they live in true sympatry or if they inhabit separate biotops in the same locality.Unfortunately, no specimens of P. caucasia or P. microlepis from Tadjikistan were represented in the study made by BaiG (1992) and also ananJeva & orlova (1979) did not give any detailed account of these populations.It appears that P. caucasia and P. microlepis are sympatric in some parts of Turkmenia and northeastern Afghanistan.This is one of the reasons not to consider them as taxa of the same species, despite the fact that they share some common characteristics.
Habitat.The Caucasian agama lives on rocks, screes, and clay slopes up to 3000 m elevation (trentev & Chernov 1949).ClarK & ClarK (1973) also confirmed its existence in mountain habitats and its absence from open plains.They further stated that it is so habitat specific that the exact limit of its range could be accurately predicted with the change of habitat.
Remarks.The lizards are known to hibernate in large groups of several hundred individuals.They mainly feed on insects when they are abundant in spring and early summer, but in autumn they switch to feeding more on vegetation (niKolsKy 1915).
The recent study on the P. caucasia complex made by Panov & zaKova (1995) and an earlier work by Panov et. al. (1987) present an intriguing hypothesis.They believe that the northernly distributed P. cauca sia and the southernly distributed P. microlepis have reached the species level of divergence.In the zones of secondary contact interbreeding takes place among them.They identified different population groups in this area and believed that populations from lower Sumbar Valley and Meshad Sands were of hybrid origin.They also observed a high level of divergence among different populations of P. caucasia.
Growth pattern of the Caucasian agama has been studied by zaKova & Panov (1991).They determined the age of individuals with sizes up to 92 mm with the accuracy of up to one year, those with sizes of up sion; yellow ocelli or transversely expanded yellow spots with black border on the lateral sides of body with black specklings, pattern of yellow ocelli or spots sometime not distinct and black spots form a network; gular region reticulated but not necessarily extensively and in some cases only few bars can be observed mainly towards tip; belly pale yellow and sometimes with black spots, breeding males have bluish grey wash ventrally.
Taxonomy.annanJeva & orlova (1979) compared different populations of P. caucasia using several morphological characteristics and suggested a splitting of the species into eastern and western groups.Unfortunately, they did not include a population from Tadjikistan in their comparison.However, they identified two populations from Turkmenia where one is comparatively longer and with higher scale counts as the other one.Later on, ananJeva & ataev (1984) described a new subspecies as Stellio caucasius trian nulatus from Turkmenia and distinguished it from the nominate form by the forementioned characters and in addition by up to three whorls per tail segment.BaiG (1992) placed this subspecies to P. microlepis, because of the higher scale counts (a diagnostic character of P. microlepis) and the relatively larger size in comparison to P. caucasia.Nevertheless, he (BaiG 1992) also noted that further studies and molecular analysis are required to support this taxonomic conclusion.And later, MaCey et al. (1998) recognized P. microlepis and P. caucasia as different evolutionary lines.ananJeva & KalyaBina-hauF (2006) analysed the relationships between P. caucasia and P. microlepis on the basis of molecular and morphological data and placed the subspecies triannulatus in synonymy of P. caucasia.However, we suggest that the forementioned population of Turkmenia with long body length and high scale counts together with the subspecies described by ananJeva & ataev (1984) should be both included in P. microlepis.
Distribution.Transcaucasia, Turkey, northern Iran, northern Afghanistan, Turkmenia, Uzbekistan, and Tadjikistan.Khan (2002) also recognized the species from Waziristan and northern Baluchistan in Pakistan.Southern parts of Iran, Pakistan and Afghanistan are occupied by P. microlepis.
Because of its wide range, P. caucasia is probably the most studied taxon among its genus.niKolsKy (1915), with reference to previous publications and museums records, mentioned several localities including Transcaspian Territory, Baku, Tbilisi, Kirovabad, Araks, Sevan Lake, Lenkoran, Khurasan and Kopet Dagh.annanJeva & orlova (1979) thoroughly reviewed this species and added two new territorial records, the Black Sea coast of Caucasus and southern vertical series but constitute a band; several enlarged mucronate scales on flanks may constitute a patch on the flanks; dorsolateral fold marked with groups of enlarged, spinose scales along entire length; other small dorsals distinctly smaller than enlarged ones; ventral scales sometimes smooth but usually carinated, smaller than enlarged vertebral; gular scales smooth but mucronate; total number of scales around midbody 86 -94 (89.4 ± 3.1); limbs strong, covered with enlarged mucronate scales, hind-limb roughly equal to or greater than the distance between gular fold and cloaca; fingers and toes compressed, lamellae 15 -17 (16 ± 0.53) under 3rd finger and 23 -25 (23.9 ± 0.83) under 4th toe.Tail depressed, oval in cross section; tail segment distinct, each segment consists of two whorls of enlarged mucronate scales; 24 -29 (27 ± 2.0) scales in first complete whorl around the tail; 3 -5 rows of callous glands present in males at precloacal and large patch at abdominal position, also represented in females (not always) but only at precloacal position.
Colour pale brown above with irregular dark brown scales which are sometimes arranged in transverse series; head yellow and may be speckled with black; transverse stripes on the upper parts of legs and tail; under parts yellow, gular may show black marmoreal pattern but breeding males may be almost black ventrally; in life displays orange shade ventrally.
Taxonomy.tuniyev et al. (1991) described the subspecies P. erythrogaster nurgeldievi but its validity was questionable and it is herein placed in the synonymy of the nominate form.The description is based on morphological differences, but MaCey et al. (1998) did not found any considerable differences using mtDNA and also recognized it as synonym.The nomen 'gaster' is a noun and not flectible.Therefore, the name 'erythrogaster' stays as masculine noun also with feminine Laudakia.to 116 mm to two years and the bigger ones with the accuracy of up to three years or more.They believe the lizards continue growing during their whole life which may be 12 -13 years.Panov & zaKova (1995) studied also the social organization and demography in this agama.They found that populations were stable with low turnover.They attribute delayed reproduction, longevity and a sedentary life style as the most plausible explanations for this observation.Older age classes dominated the age structure of all the subpopulations studied.They further mentioned that it is a good example of lizards practicing a K-strategy.
Paralaudakia erythrogaster (nikoLsky, 1896) Taxonomy.ananJeva et al. (1981) described a new species as Agama chernovi from Tadjikistan.They compared the species with P. himalayana and P. badak shana and found several dissimilarities from the forementioned species.Unfortunately, they did not compare their specimens with Agama isozona, which was also described from the same area before.An examination of the type specimen of the latter taxon, deposited at the Natural History Museum in Vienna, shows marked differences to P. himalayana, to which it was synonimised by sMith (1935).Simultaneously, it has and Himalaya up to Tibet.Khan (2002) documented the species from Gilgit and Chitral in the remote areas of northern Pakistan at elevations between 3000 and 3200 m.All the three forementioned species are distributed at an elevation range of about 1000 -3500 m a.s.l.Pa ra laudakia (himalayana) himalayana mostly remains over 2000 m.BaiG (1992) showed doubts about the type locality of P. badakshana (Mazir-i-Sharif, NW-Afghanistan), but however the localities mentioned by anderson & leviton (1969) for the paratype series are very likely.The suspicions about Maziri-Sharif as type locality are mainly because i) the loca lity is far from the main distribution area of this group, and ii) the elevation of the area (about 360 m a.s.l.) is unbelievable for either of P. badakshana, P. bo cha riensis or P. himalayana.
Habitat.BaiG (1992) collected several specimens of P. himalayana in the northern mountain areas of Pakistan where cliffs and boulders are avoided.It was found running on the sandy bank of a river or away from the riverside among small stones which are used for hiding.The lizards were taking shelter in the crevices of stone walls along the river or road-side, or underneath larger stones or slabs of stones.Their larger number along the watercourse may be related with the availability of food, otherwise they are not found close to this habitat.

Remarks.
The distribution ranges inhabited by the taxa of the superspecies complex apparently lie quite close to each other within an intricate system of mountain ranges.These ranges sometimes attain heights well above 5000 m a.s.l.Several peaks remain under snow throughout most of the year, forming impassable barriers for these lizards and dividing them into many disjunct populations.It appears that speciation of this group occurred at Pamir Knot where Pamir, Karakoram, Hindu Kush and Himalaya meet each other.Paralaudakia himalayana went into the eastern side and is presently distributed in southern Pamir, Wakhan, and mountain ranges of Karakoram, Ladakh and Himalaya up to Tibet.P. bochariensis followed a northwestern route and is presently distributed at different biotops on the western side of Pamir in Tadjikistan and Uzbekistan.The southwestern side along the Hindu Kush range was followed by P. badakshana which is now found in Badakshan, Kabul and Ghazni provinces of Afghanistan. ananJeva et al. (1981) observed P. himalayana at one locality of Zeravshan mountain range.BaiG (1992) also came across with two specimens (ZFMK 32157 -58) from Schador (Tadjikistan).All remaining records from Tadjikistan and Uzbakistan do not show P. himalayana in the west or northwest of Pamir.The very small, smooth; skin of lateral sides loose forming dorsolateral fold; total number of scales around midbody 122 -166 (132 ± 8.35); limbs less strong, covered with enlarged mucronate scales, hindlimb little longer than distance between gular fold and cloaca; fingers and toes compressed, 16 -23 (19.7 ± 1.73) lamellae under third finger and 24 -32 (27.3 ± 1.93) under fourth toe, fourth toe considerably longer than third, extremity of the claw of the latter does not reach the base of the claw of former.Tail moderately depressed, oval in cross section; each tail segment (indistinct) consists of three complete whorls of moderately enlarged spinose scales; 26 -43 (35 ± 5.0) scales in first complete whorl around the tail; callous glands present in males only at precloacal position, number of rows at precloacal position never exceed three.
This species exhibits sexual dimorphism in color pattern, which is more sharply demarcated in females as compared to males.Females have silver grey or olive ground color versus a dull brown or yellowish brown in males.Head shows dark spots over light ground; gular reticulated; chest and belly pale yellow in females and juveniles but is blue or a grey wash in males; vertebral stripe grey, irregular black spots form festooned band on each side of vertebral line; olive grey above with dark rimmed yellow ocelli along the entire vertebral length in females, in males these ocelli are mainly concentrated on neck region; tail with dark spots which sometimes gives the impression of cross bars, the banded pattern is usually seen in females or juveniles.Females are clearly distinguishable from males by the presence of a red or bright orange collar region; juveniles have this female pattern.
Taxonomy.Together with P. bochariensis and P. badakshana this species constitutes a sub-group within Paralaudakia and is being recognized as supraspecies complex.The group may be characterized as small Paralaudakia lizards whose SVL rarely exceeds 90 mm.However, ananJeva et al. (1981) reported the range of A. chernovi (recognized as synonym of P. bochariensis) as 80 -120 mm, but without presenting information about the measured specimens.Therefore, BaiG (1992) studied several specimens of A. chernovi including a type specimen and none were found longer than 90.00 mm (with the exception of ZMB 46603 which was 96.00 mm).There are also some few specimens of P. badakshana from Kabul, Afghanistan which have a range of 90 -92 mm, but all other examined specimens belonging to this group were measured with a SVL of 90 mm or below.In particular, P. himalayana does not even reach 90 mm and females are even shorter than males.

Distribution. Southern Pamir, Wakhan Corridor in
Afghanistan, mountain ranges of Karakoram, Ladakh praoccular scales which are smooth, at anterior part of head at the level of anterior margin of eye flower shape formation is quite marked; labials 11 -15 (13 ± 1.0); groups of highly spinose scales present on neck and sides of head, preauricle constitutes circular series; vertebral scales heterogeneous, strongly keeled, vertical series of distinctly enlarged scales with distinct ridges and spines interrupted by other moderately enlarged scales; several enlarged mucronate scales randomly present; dorsolateral fold marked with enlarged, spinose scales along entire length; other small dorsals distinctly smaller than enlarged ones; ventral scales smooth, smaller than enlarged dorsal; gular scales smooth; total number of scales around midbody 88 -109 (98 ± 5.0); limbs strong, covered with enlarged mucronate scales, hind-limb about equal or more than the distance between gular fold and cloaca; fingers and toes compressed, lamellae 14 -20 (16.4 ± 1.2) under 3rd finger and 20 -26 (23.7 ± 1.4) under 4th toe.Tail depressed, oval in cross section; tail segment distinct, each segment consists of three whorls of enlarged mucronate scales, near its origin mid-dorsal rows constitute two whorls in each segment; 22 -30 (26 ± 2.0) scales in first complete whorl around the tail; 3 -5 rows of callous glands present in males at precloacal but not at abdominal position; not represented in females.Colour olive yellow or olive grey above with irregular black scales; head pale brown or grey; upper parts of legs and tail usually speckled with black but sometimes may show banded pattern; under parts yellow usually spotted with black; throat in life shows black & orange spots which may appear or disappear.
Colour pattern of live specimens according to terent 'ev & Chervov (1949) Colour is highly variable in different populations.Generally olivaceous above with vertebral zone usually yellow, sometimes with distinct lateral extension and yellow ocelli or transversely expanded yellow spots with black border similar to P. caucasia but some of Afghan specimens are bluish grey with yellow spots arranged in transverse series, pattern of yellow ocelli or spots usually distinct in upper half; gular region may be reticulated; belly pale yellow and may sometimes have black spots; under parts of forementioned grey Afghan specimens were also grey with some yellow blotches on the throat.
Distribution.Southern Iran, southern & western Pakistan, Afghanistan and some parts of Turkmenia.
anderson (1974) thought that P. microlepis is a distinct population of P. caucasia only and assigned a subspecific status.The same author also expanded the range from the type locality to central and south eastern parts of Iran up to Afghanistan and Baluchistan.Minton (1966) and Mertens (1969) studied the herpetology of Pakistan and were unable to find P. mi crolepis.They identified the Pakistani population as P. caucasia, which is contrary to BaiG (1992) later.Although most of the material from Pakistan which is included in BaiG (1992) was identified as P. micro lepis.However, some records show that P. caucasia is present in northwestern Pakistan, while Baluchistan and southern parts of the country are occupied by P. microlepis.BaiG (1992) has extended the range of P. microlepis by including some localities in western and previously published record from Afghanistan, however they reported the presence of this species from Mazar-i-Sharif (Afghanistan) at an elevation of 457 m.The material studied by BaiG (1992) shows its presence in Badakshan (NE-Afghanistan) from an elevation of 1600 m.
Habitat.According to terent'ev &Chernov (1949) andananJeva et al. (1998) it is a mountain lizard, present at an altitude up to 2600 m above sea level.It can be found on rocks and stones of steppes, slopes of canyons, and on more or less gentle slopes covered with coarse disintegrated rock formations.Additionally, sometimes near mountain rivers and creeks.In some places they have also been found near mountains in loessy hills.The space underneath or between stones, or the fissures in rocks or slopes serve as their hideouts.
Remarks.Eggs were laid not earlier than end of June, or the first half of July.One female lays two to three eggs of 18 -20 mm in length.At altitude of 1500 m a.s.l.young start appearing at the end of September.This species feeds on various insects and arachnids.
Paralaudakia microlepis (BLanForD, 1874)  The colouration of this form is speckled with black on pale or lemon yellowish ground; head lemon yellow with several black spots; yellowish lateral extensions from vertebral line more distinct in anterior half which itself is darker than posterior part; gular dark grey with yellow spots or reticulated in juveniles; belly pale yellow with black spots; chest dark grey; tail light in proximal part and shows indistinct banded pattern and is dark distally.Peters (1971) is mentioned as author of this subspecies.However, he mentioned himself that MunKhBayar & shaGdarsuren (1970) described the same taxon under the same name some months before and therefore, Peters (1971) created a primary homonym.The original description by MunKhBayar & shaGdarsuren (1970) is based on a holotype (no.611) and 30 paratypes (fig.6), all collected in the area of the source of Uliastajn [=Uliastai] River (Khovd district) from 1964 -1970 (see below).The holotype (no.611) was originally deposited at the herpetological laboratory of the "State University for Education of Mongolia".Later it was transferred to the University of Tashkent in 1970 and is today most probably lost (pers.com.Kh.MunKhBayar, March 2011).Moreover, there are two paratypes in the latter mentioned collection, one paratype in the collection of the "Zoological Museum of the Mongolian State University" and three paratypes in the biological collection of "School numer 33".All remaining paratypes are still at the collection of the "State University for Education of Mongolia".Currently, one of these southwestern Pakistan, almost all of Afghanistan (at several places together with P. caucasia), and southwestern Turkmenia.

Taxonomy. In all recent publications
Remarks.Paralaudakia microlepis was largely neglected or less attended by herpetologists in the past.It was described in 1874 by BlanFord from southern Iran, north of Shiraz.In subsequent publications it was accordingly mentioned, but records were largely based on the original description, except for ClarK et al. (1969).Panov et al. (1987) and Panov & zyKova (1995) studied populations of rock agamas from Trans caucasia, Middle Asia, Iran and adjacent regions and mentioned that populations (especially in and around SW Turkmenia) showed marked differences.They suspect that some of them are a product of secondary contact between P. caucasia and P. microlepis because they exhibit hybrid characteristics.
Diagnosis.Differs from the nominate form mainly by having tail segment of 3 whorls; slightly less number of scales around tail and body; scales generally more carinated; relatively shorter hind-limbs; callous glands present in males only at the precloacal position but rarely females also display this character.
Description.Head and body depressed, covered generally with small scales; snout longer than the distance between eye-tympanum and of eye width; tympanum exposed, in adults usually half or less than half of eye width; nostril pierced below canthus rostralis, less than half of nasal, separated by 1 or 2 scales from rostral, directing outward and backward; no gular pouch, gular plicate; upper head scales heterogeneous, smooth; labials 10 -14 (12 ± 1.0); groups of spinose scales present on the neck and sides of head especially around   The colouration of this species is generally similar to P. caucasia.Olive yellow above; head and body speckled with black, may have pale yellow ocelli especially in anterior half; gular yellow or in males grey with yellow spots; belly pale yellow but males have grey wash; tail light in proximal part and may show banded pattern whereas it is dark distally.
Taxonomy.Although this species was described in 1875 from central Asia it has attracted very little attention from herpetologists.zuGMayer (1909) very superficially described a new species as Agama tari mensis from Mongolia and both taxa were mentioned in subsequent literature until 1971.At this time Peters (1971) compared P. stoliczkana and A. tarimensis and carried out a morphological analysis of the material from Central Asia.He divided the material into four groups according to the different geographical localities viz.Kashgar, East Tien-Shan, Gobi-Altai and Mongolian Altai.
Regarding colouration Peters (1971) found no differences within populations of P. stoliczkana and A. tarimensis from the western side among the Kashgar and Tien-Shan.However, he did find differences within Mongolian populations which have a dark chin, throat, neck and chest with light spots.Moreover, he noticed that males show a more contrasting pattern by having more melanin as compared to females and secondly, northern Mongolian lizards exhibit more in-paratypes was gratefully donated to the authors by Kh.MunKhBayar and is now present in collection of the Museum Koenig in Bonn (ZFMK 93007).The authors described the holotype as an adult specimen with a SVL of 113mm and a tail length of 181mm.The tail is arranged in whorls of three scale rings each, having 121 scale rings together.
One year later, MunKhBayar (1971) placed altaica as subspecies to P. stoliczkana.This was in accordance with Peters (1971) who identified the taxa as distinct subspecies, but published his description about a year after the original description.Peters (1971) compared populations of P. stoliczkana from four different localities of Central Asia and found populations of Gobi-Altai and Mongolian-Altai as different.He distinguished his P. s. altaica by having a shorter tail and hind limb as compared to the nominate form.These characters have been later corroborated by BaiG (1992).
Distribution.Mongolian and Gobi Altai area.Mun Kh-Bayar (1971) mentioned some detailed localities within the Altai area: Tsagaan Burgas, Shar Khuls, Atas, Ya.Eglon at Altan Uul mountain, at river Bulgan Gol, Uushgiin Ulaan, Aj Bogd, Takhiin Shar Khuruu, Atas, Nemegt tost and in the Edren mountains.MunKhBayar & shaGdarsuren (1970) mentioned the river Uliastain in their description.This subspecies not only marks the northern limit of the species, but also the northeastern end of the genus Paralaudakia and extends its range into Mongolia.
Paralaudakia stoliczkana stoliczkana (BLanForD, 1875)  3) around middle of the body; 8 -10 rows of enlarged, strongly keeled vertebral scales with sharp lifted margin, median two rows may be of relatively small scales; a distinct patch of enlarged mucronate scales on flanks; other small dorsals distinctly smaller than enlarged ones; ventral scales smooth, smaller than enlarged vertebral and flank scales but larger than gular and other small dorsal scales.Limbs moderately strong, hind-limb longer than distance between gular fold and cloaca; fingers and toes long and compressed, 18 -24 (22.2 ± 1.59) lamellae under third finger and 26 -32 (29.5 ± 1.8) under fourth toe.Tail depressed, oval in cross section, forming distinct annuli, each tail segment consists of three complete whorls; 28 -33 (30 ± 2) scales in distal whorl of first proximal segment of the tail, scales on dorsal and lateral sides strongly mucronate.Callous glands present in both sexes, 3 -4 rows at precloacal and small patch at mid abdominal position in males and similar patches in females where the number does not go beyond 3 at precloacal position and abdominal patch less often present.Olive grey, variegated, spotted with yellow, juveniles and young with dark and light longitudinal stripes which may be broken, central light stripe continues on to the tail; throat with dark reticulation; belly yellowish white; tail generally light in proximal half and dark distally and shows light and dark transverse bands especially in young specimens.Rarerly exceeds 100 mm in SVL but never goes beyond 110 mm.

Distribution. Northern Punjab and northwestern
Frontier Province in Pakistan, Kashmir (at least the part now associated with Pakistan) and Afghanistan.
In Pakistan and Kashmir it occurs in the Lesser Himalayas at a low elevation of approximately 700 -1300 m a.s.l.Although there is only one record from Afghanistan, but its presence in Arandu (Chitral, Pakistan) within a mountain range which spread into Afghanistan suggest the presence of the species also in eastern parts of this country.There is an interesting pattern of the vertical distribution together with L. tu berculata in Pakistan and Kashmir.In these mountain areas L. agrorensis and L. tuberculata are distributed tensive pattern than southern (Kashgar and Tien-Shan) ones.The light ground colour in the lizards has different shades and is rather individual or locality-oriented.The yellowish green spots of A. tarimensis mentioned by zuGMayer (1909) were not seen by Peters (1971).He observed no difference in colour pattern between P. stoliczkana and A. tarimensis, but found differences in the number of scales at different body parts.However, he attributed these differences to be clinal and suggested A. tarimensis as junior synonym of P. stoliczkana.

Type species. Agama tuberculata hardWiCKe & Gray, 1827
Diagnosis.Head and body depressed; head scales smooth or rugose; no gular pouch; usually no patch of enlarged or spinose scales on flanks (but present in some species); gular scales smooth; ventral scales smooth; vertebral scales enlarged, sometimes with median rows of small scales; groups of spinose scales on neck and sides of head; tail segment of two to four whorls, usually three to four.
Description.Head very depressed; snout longer than the distance between eye-tympanum; tympanum ex-scales keeled, larger than other small dorsals, 10 -12 oblique rows of strongly keeled scales; patch of enlarged mucronate scales on flanks and similar scales scattered over the sides of body; other small dorsals distinctly smaller than enlarged ones; ventral scales smooth, smaller than enlarged vertebral and flank scales but larger than gular and other small dorsal scales; gular scales granular, smooth; skin of lateral sides loose forming a dorsolateral fold; total number of scales around midbody 102 -130 (118 ± 8.43); limbs strong, covered with enlarged mucronate scales, hind-limb slightly longer than distance between gular fold and cloaca; fingers and toes compressed, 20 -30 (26.5 ± 2.1) lamellae under 3rd finger and 27 -37 (34.9 ± 2.2) under 4th toe.Tail depressed, oval in cross section; each tail segment consists of 4 whorls of enlarged mucronate scales with three remaining on the ventral side; 26 -33 (29 ± 2.0) scales in first complete whorl around the tail; callous glands present in males at precloacal and also at abdominal position, number of rows at precloacal position 4 -6 and an oblong patch on abdomen; very rarely represented in females at precloacal position in three or less rows.Head pale yellow; gular spotted or speckled with black, males marked with blue, in juveniles reticulated; chest and belly pale yellow but may be speckled with age, males with blue or grey wash; numerous yellow spots on dark slate background, sub-adults have irregular pattern; tail light in proximal half and dark distally, young or juveniles sometimes show faded pattern.
Taxonomy.BaiG (1992) studied several specimens from different localities, including typematerial of L. tuberculata, L. sacra, B. sykesii and L. dayana, and found several dissimilarities among specimens from western mountain ranges.However, on the eastern side of their distribution, up to Sikkim, all material is more or less similar to L. tuberculata.
The types of B. sykesii are dissimilar.BMNH 1946.8.28.16 from Ladakh belongs to L. dayana, whereas BMNH 1946.8.28.60 (without locality) was identified as L. tuberculata.The types of L. dayana and few more specimens from the same locality are clearly distinct from L. tuberculata.They show a strong carination, quite distinct tail segments and patches on the flanks.Therefore, as recognized by BaiG (1992), L. dayana should be recognized on species level, while B. sykesii can partly be identified as L. tuberculata and L. dayana.
In Simla, L. dayana occurs at an elevation of 3000 m, while L. agrorensis, if allopatrically distributed, occurs at lower elevations of about 1300 m a.s.l.parallell, with L. agrorensis at low (700 -1300 m) and L. tuberculata at relatively high (1200 -2200 m) elevation.These upper and lower limits vary slightly in different areas within their specified ranges, but occasionally they are found in sympatry.L. agrorensis may also be found in sympatry with another agamid, Calotes versicolor in Pakistan.
Habitat.Laudakia agrorensis is a mountainous species and occurs in low mountain areas with vegetation cover.Several individuals were found basking or running on rocks, mostly along water streams.Availability of food (insects) is the most probable reason for their presence in that habitat.Instead of single or solitary pairs, large groups of individuals were usually seen.Except the studies of callous glands (BaiG & BöhMe 1991b) and some morphological comparisons, other aspects of this species are largely unexplored.
Remarks.Among the specimens housed in different museums, BMNH 1933.4.1.39differs from L. agro rensis in several characteristics which are diagnostic for L. pakistanica and therefore the specimen is reidentified as the latter species.Diagnosis.Head and body depressed; head scales carinated; patch of enlarged, mucronate scales on flanks, similar scattered scales on body; groups of low spinose scales on neck and sides of head; tail segment of four whorls dorsaly becoming three ventraly; callous glands in males, very rarely occurring in females.
Description.Head and body depressed; snout more than one and half times that of the distance between eye-tympanum or eye width and more than twice the tympanum diameter; tympanum exposed, more than half of eye width; nostril pierced on or below canthus rostralis, less than half of nasal, pointing outward and backward, touching rostral; no gular pouch, gular plicate; upper head scales heterogeneous, subequal, carinated; upper labials 9 -13 (11 ± 1), lower labials 9 -11 (10 ± 1.0); groups of low spinose scales present on the neck and sides of head especially around tympanum, preauricle in series otherwise in groups; vertebral Vertebrate Zoology n 62 (2) 2012 total number of scales around midbody 120 -156 (137 ± 13.4); limbs strong, covered with enlarged mucronate scales, hind-limb usually greater than the distance between gular fold and cloaca; fingers and toes compressed, lamellae 17 -19 (17.8 ± 1.0) under 3rd finger and 22 -27 (24.3 ± 1.8) under 4th toe.Tail moderately depressed, oval in cross section, distally rounded, much longer, usually 2.5 times or greater than the distance between snout to vent; tail segment becomes distinct slightly distal to its origin, not consistent in number of whorls in each segment, 2 -3 whorls in each segment but gradually changes into five (sometimes six) in terminal part of tail; mucronate scales in proximal part of tail have very low ridges; 18 -28 (21 ± 5.0) scales in first complete whorl around the tail; 4 or 5 rows of callous glands present in males at precloacal and a patch (sometimes small and rounded like P. nupta and sometimes broad and rectangular) at abdominal position.Colour of this species is highly variable.Those in Salt Range have light brown head with reddish tinge on posterior side; rest of the body olive brown with several yellow spots all over the body; under parts pale yellow, gular reticulated; tail light brown in proximal half and black in distal half.sMith (1935) mentioned five specimens collected at Ladha (Waziristan) by CaPt.inGoldBy as dark brown all over dorsally.According to stoliCKzKa (1872) juveniles are olive above, yellowish white below; entire head, including chin and front breast, reticulated with black; neck, body, limbs and base of tail above with numerous small black and interspread yellow spots; eyelids and supracilliary ridge yellow; tail dusky black toward tip.The adults are more brownish olive, with dark reticulation on the upper head less distinct, the black spots on the body small and more or less confluent, but the yellow spots are more brightly coloured and larger in size; tail pale yellowish at the base but for the greater part of length entirely black.In spring males are more or less jet black, usually on the tail and posterior portion of the back (BlanFord 1876).Minton (1966) mentioned that adult males are sooty black above and black to dark grey beneath; adult females dark brownish grey with traces of paler dorsal spots, light grey to dull white tail at the base and underside of thighs; juveniles dusky grey, with faint dorsal spots, belly white, throat mottled with grey.He also observed colour differences among members of the same sex but of different age groups in Ormara (Mekran Coast).
In general this species resembles L. nupta, and in some localities of southern Pakistan both of them are found together.However, they are distinguished by several morphological characteristics.Laudakia species have a tendency to change the number of whorls in caudal segments from the anterior to posterior region, but the difference never exceeds one whorl and Nothing is known about the vertical distribution pattern between L. tuberculata and L. dayana in Kashmir (the part presently under the administrative control of India).Future studies proving allopatric distributions of these two species may identify them as two distinct populations of the same species only, but at the moment both are recognized as full species because of distinct morphological characters.

Remarks.
In scalation and colour pattern this species also resembles L. agrorensis, which is relatively smaller, possess three whorls per tail segments and shows a different arrangement of callous glands.Color is grayish brown with yellow spots all over the body, including head but excluding tail.Under parts yellowish gray with yellow speckling, which are prominent at chest, gular region and chin.Proximal part of the tail yellowish gray, distal dark brown.
Laudakia nupta fusca (BLanForD, 1876)  it usually occurs only on the dorsal part of the tail.Accordingly, this species, because of its inconsistency of tail segments, is unique and distinguishable from all other members of the previsouly collective genus Laudakia.Secondly, the callous abdominal patch is usually oblong or a thin longitudinal stripe in Paralaudakia or Laudakia, but it is rectangular, much wider than long, in some specimens of L. melanura (BMNH 76.10.13.1,98.6.29.1 and PMNH 368).
Taxonomy.BlanFord (1876)  Description.Head quite heavy, only moderately depressed, subtriangular when viewed from above; body not too depressed but rather slender; snout longer than the distance between eye-tympanum, about twice that of eye width; tympanum exposed, large, equal or more than that of eye width; nostril pierced on canthus rostralis, more than half of nasal, separated by two scales from rostral, directing outward; neck narrow; no gular pouch, gular plicate; upper head scales heterogeneous, subequal, mostly smooth, on posterior margin rugose; labials 12 -17 (15 ± 1.0); groups of highly spinose scales present on neck and sides of head especially around tympanum; vertebral scales strongly keeled, distinctly enlarged and constitute a very broad band of 16 -18 scales which cover most parts of the dorsal body and resemble a flask, narrow near neck and broad posteriorly; no patch of enlarged mucronate scales on flanks; other small dorsals distinctly smaller than enlarged vertebral; ventral scales smooth, smaller than enlarged vertebral; gular scales about the size of ventrals, smooth; total number of scales around midbody usually 75 -106 (92 ± 9.09); limbs strong, covered with enlarged mucronate scales, hind-limb usually greater than the distance between gular fold and cloaca; fingers and toes compressed, lamellae 15 -18 (16.8 ± 0,9) under 3rd finger and 19 -24 (21.3 ± 1.5) under 4th toe.Tail moderately depressed, oval in cross section, distally rounded; first tail segment distinct, only little away from pelvic, each segment consists of three whorls of enlarged mucronate scales but occasionally may be four in terminal end; 16 -29 (23 ± 3.0) scales in first complete whorl around the tail; 3 or 4 rows of callous glands present in males at precloacal position and a small patch at abdominal position, may be represented in females but very rarely.
Colouration is predominately uniform yellowish, reddish brown or may be speckled; head dark brown or blue in males; tail yellowish brown in proximal half Diagnosis.Size large; head heavy; nostril on canthus rostralis; body moderately depressed; absence of a nuchal fold across the neck; vertebral enlarged scales strongly mucronate and cover most of dorsal part of body; no patch of enlarged mucronate scales on flanks; tail long and has segments of three whorls; callous glands present mostly in males.
Description.This taxon differs from the nominate form by some slightly expressed characters, and the colouration of adult males.The striking character is the absence (sometimes rudimentary obvious) of the nuchal fold across the neck.It usually has less than 90 scale rows along the midbody, but other characters are often shared by both taxa and only lower or stronger expressed in this subspecies: tufts of spiny scales around the ear and on the sides of the neck large and more numerous, enlarged mid-dorsal scales (in 13 -16 rows) more strongly mucronate, and scales of base of tail much larger than in the nominate form.
Adult males with an entirely carany-yellow head, especially during the breeding season; body, limbs and tail dark brown to black above; belly, underside of legs and tail black sparsly speckled yellowish; callose ventral and precloacal scales amber.Females brown, speckled yellowish; distal part of the tail black.Juveniles more similar to females than to males; with a greyish, yellow body colouration with an irregular pattern of brownish crossbars or reticulated pattern; head black, often with yellow chevrons and yellow spots on the upper head; tail banded with a black tip.
Distribution.The subspecies is only known from south eastern Iran, Pakistan.It was collected in the Ba lu chistan Province at Kalagan and near Jalk by Blan Ford (1876).Minton (1966) recognized it from southern and southwestern Pakistan (near Khadeji Falls, 30 miles NE of Karachi; near Diwana on the upper Hab River; southern part of Pab Hills).Khan & Mirza (1977) also found it in southwestern Ba luchistan.anderson (1999) mentioned a specimen from Binak at the Persian Gulf coast also belonging to L. n. fusca.According to Minton (1966) the subspecies is knowm from elevations up to 2000 m a.s.l.

Habitat.
A shy and secretive lizard, only found in rocky areas with vertical or nearly vertical surfaces that provide shelter.According to rasteGar-Pouyani & nilson (2002) also the type of vegetation has an effect on the distribution range of the subspecies.ings) indicate that the area of activity of an individual generally has a radius of less than fifteen meters.The basking area, as indicated by the distribution of fecal pellets, is usually not more than two to three metres (anderson 1963).
anderson (1963) noticed shift in the periods of activity from February through the summer.In late February and March individuals were observed only at midday and early afternoon while basking on rocks and exposing maximum surface area to the direct sunlight during air temperetures of about 30 °C.In the afternoon they were raised on their forelegs, facing the sun that the sunlight strikes the head and belly, and the angle of reflection from the surface of the rock exposed them to maximum radiation.Later spring, basking was initiated progressively earlier in the day and continued later into the afternoon.In summer the lizards retired during the hottest hours, and their activity was restricted to the earliest daylight hours and late afternoon in July and August.Additionally, anderson (1963) measured a cloacal temperature of 27.2 °C of a basking individual during an air temperature of 22.4 °C.The critical maximum temperature in this species was recognized between 43.8 -48.5 °C.
Analysis of stomach contents shows both insects (mostly beetles) and plant material.Females containing gravid eggs were collected in March, August and October, indicating that eggs are laid at least in the spring and in the autumn in Iran region (anderson 1963).
Laudakia nuristanica (anDerson & Leviton, 1969) 1969 Agama nuristanica anderson & leviton,Proc. Calif. Acad. Sci.,Fig. 8. Type locality: "Kamdesh, eastern Afghanistan, 1342 metres elevation." Diagnosis.This form is very close to L. tuberculata and may be distinguished by having a higher number of scales around the body and tail, indistinct tail segment, and a unique type of scattered, enlarged conical, spinose scales over the limbs and body.Khan (2002).Although the trinomial status has been adopted by several authors, BaiG (1992) recommended the point of view adopted by sMith (1935) and found additional support from the observations made by anderson (1963).However, rasteGar-Pouyani & nilson (2002) also found good differences between the taxa and followed authors like Minton (1966) and Mertens (1969) Colouration is dark grey or brown marblings on pale brown ground colour; vertebral stripe is pale yellow; head dusty brown with dark brown specklings, three supracilliary dark bars; throat brown or silver grey with yellow ocelli which are more concentrated near tip; distal part of tail dark while proximally it is pale brown; belly and under parts pale yellow.
Taxonomy.The species is very close related to L. day ana and L. tuberculata.anderson & leviton (1969) compared it with material of L. tuberculata from Nepal and found several striking dissimilarities.BaiG (1992) suggested that it undoubtedly differs from L. tuber culata, but it is more similar in some characters (e.g.number of scales around the body, scattered enlarged scales on flanks, rugose head scales) to western populations as to Nepalese populations of L. tuberculata.In tail characters it differs both from L. tuberculata and L. dayana by having four complete whorls around the tail.The arrangement of the scales over the limbs is unique i.e. enlarged conical scales among other very small scales.

Distibution. Eastern Afghanistan and Northwestern
Pakistan.
The species was previously known only from Afghanistan, but BaiG (1988a) recognized its presence in the Chitral district of Pakistan.This mountain range streaks from Afghanistan into Pakistan therefore a Diagnosis.This form is very close to the nominate form but having fewer scales around the body and a more mosaic-like in colouration.
Description.Head triangular when viewed from above, depressed; nostrils oval, touching edge of canthus rostralis, pointing outward and backward; upper head scales heterogeneous, subequal, flat; upper labials 11 and lower labials 11 to 12; groups of spinose scales present on the sides of the head and neck, those at subauricle level arranged in two distinct horizontal rows; vertebral scales strongly mucronate, enlarged vertebrals start from postshoulder level and gradually increase from anterior to posterior, eight rows of strongly mucronate scales with sharp lifted margins, median two rows consist of relatively small scales which may overlap in posterior half of body; a distinct patch of enlarged, distantly arranged, spinose scales whose lifted margin is broken into sharp spiny border, other small scales spinose or multispinose; other small dorsals distinctly smaller than enlarged ones; ventral scales smooth, smaller than enlarged vertebral and flank scales but larger than gular and other small dorsal scales; skin of neck and lateral sides loose forming gular and dorsolateral folds respectively, posterior half of dorsolateral fold marked with enlarged, strongly mucronate scales; total number of scales around midbody 164; limbs strong, covered with strongly mucronate above and flat scales below, hindlimb longer than distance between gular fold and cloaca; 21 lamellae under third finger and 27 under fourth toe.Tail depressed, oval in cross section, each tail segment consists of three complete whorls; 34 scales in first complete whorl around the tail; scales on dorsal and lateral sides strongly mucronate and spinose, spine extends little beyond posterior margin of scale; ventral scales smooth.Callous glands present, six rows at precloacal and 2 -3 scale wide small patch at mid abdominal position.
Colouration is a black mosaic on silver grey ground, head black with scattered light colour scales; forelimbs, shank and foot show banded pattern; thigh and proximal part of tail possess scattered black scales; distal 2/3 of tail black; gular region with yellow bloches on black ground; chest, belly and lower parts of limbs pale yellow with irregular dark spots.
Distribution.In and around Chilas, Federal Ad mini stered Northern Areas, Pakistan.tral scales smooth.Callous glands present, six rows at precloacal and a large oblong patch at mid abdominal position.
Colouration is a dull dark brown (in formalin has changed into dark grey) with distinct white cross bands; head and distal part of tail black; limbs show highly indistinct pattern; gular region bluish black with irregular pale yellow bloches; chest, belly and under parts of limbs pale yellow with thick black reticulation.
Laudakia pakistanica was previously considered to be restricted to a small radius of about 40 km around Gilgit, Pakistan (BaiG 1989), but the description of two new subspecies considerably enlarged the distribution range.BaiG (1992) suggests that L. pakistanica is widely distributed along Indus from Gilgit down to the Mansehra district of Pakistan.The previously defined range is occupied only by the nominate form and the subsequently defined range extensions are inhabited by different populations of L. pakistanica (BaiG & BöhMe 1991a).
Habitat.It is restricted to montane regions and can easily observed on cliffs or while basking on rocks during daytime.The habitat of all three taxa within L. pakistanica is typified by barren rocky mountain with very sparse vegetation.However, the southern limit contains more green eco-zones, but the species itself remains restricted to unvegetated rocks.More frequent presence is generally witnessed along the water course.
Remarks.In early afternoon, when temperature rises up to or above 40°C individuals were often found on the shaded side of the rocks.Even that the mentioned habitat are rocky areas with sparse vegetation, individuals can be observed in vegetated area or on river or stream banks, presumably for insect hunting.Analysis of the gut contents shows that it is an omnivorous species, feeding on insects (mostly beetles) and herbs (BaiG 1989).
BaiG (1989) described it as a non gregarious species, always found them solitary or in pairs, which is supported by the presence of callous glands in females as reported by BaiG & BöhMe (1991).The subspecies appears unique among all members of Laudakia in having 100% callosity irrespective of sex, and provides indirect evidence linking territorial behavior and callosity.However, further studies are still required to support or disprove this hypothesis.
Distribution.In a radius of about 40 -50 km around Gilgit (Federal Administered Northern Areas of Pakistan).
Laudakia papenfussi zHao, 1998 Callous glands present, six rows at precloacal and 2 -3 scale wide small patch at mid abdominal position in males and similar patches present in females but does not extend beyond three rows at precloacal position and rarely possess an abdominal patch.
Colouration is a jet black above; under parts speckled with yellow or orange.However, young and juvenile specimens may show some faded pattern above and less black underneath.This indicates that colour variations in the nominate form are somehow related with age.
(192 ± 6.0) (according to ananJeva et al. 1990, up to 239 scales); limbs strong, covered with enlarged scales, over thighs mostly smooth and weakly keeled except those on the posterior border of thigh, hindlimb little longer than distance between gular fold and cloaca; fingers and toes compressed, usually 21 -27 (23.6 ± 1.99) lamellae under 3rd finger and 28 -32 (30 ± 1.4) under 4th toe.Tail depressed, oval in cross section; each tail segment consists of 4 whorls of enlarged mucronate scales but only 3 on the ventral side, not marked in proximal part; 30 -32 (31 ± 1.0) scales in first complete whorl around the tail; callous glands present in males at precloacal and also at abdominal position, number of rows at precloacal position 4 -6 and an oblong patch on abdomen.
Colouration in preserved specimens is a dull grey or brownish speckled with dark spots forming a very indistinct pattern; gular spotted or reticulated; tail light in proximal half and dark distally.
Taxonomy.On the basis of i) the much larger size, and ii) the presence of large patches of callous scales at precloacal and abdominal positions, sMith (1935) described L. sacra as a subspecies of L. himalayana.All subsequent authors (e.g. anderson & leviton 1969(e.g. anderson & leviton , ananJeva et al. 1981) followed this taxonomic conclusion and recognized L. sacra as subspecies.However, ananJeva et al. (1990) realized the complexity of the situation, found striking dissimilarities between L. sa cra and L. himalayana, and raised it to full species level.Unfortunately, they compared it extensively with L. himalayana, but failed to include L. tuberculata in their comparison.Although some differences are obvious, BaiG (1992) suggests that it should be placed in the L. tuberculata group because of the similarity in major characteristics (e.g.size, number of scales around body, tail segment, callosity).This is recently supported by MaCey et al. (2000) who recognized L. sacra as the sister taxon to a clade containing L. nupta and L. tuberculata.

Distribution.
Restricted to the river drainage of the Yarlung Zangbo in Lhasa Valley, Xizang (Tibet), Autonomous Region, between elevations of 3000 -4000 m a.s.l.
Previously it was only known from the type locality, but ananJeva et al . (1990) recognized it from several other localities and extended the range to Bomi (29°50'N, 95°45'E).The distribution range lies quite close to the eastern limit of L. tuberculata and includes an area where both species are allopatric.Laudakia sa cra inhabits an isolated area in southeastern Tibet.The elevational range is given by ananJeva et al. (1990) as 3000 -4000 m a.s.l., which is apparently the highest among all Laudakia species and the species also marks the southeasternmost limit of the genus.vertebrals; limbs stout, longest toe of hind limb reaches the tympanum, back of limbs covered with large, strongly keeled scales, scales of inner side of upper arm small, fingers and toes well developed, claws compressed and sharp.Tail cylindrical, slightly depressed at the base, covered with large, strongly keeled scales, scales of lower surface largest, scales are arranged in segments, each segment with three rings of scales.
Distribution.Only known from the type locality (Mayang River Valley between Mayang Village and Diya Village, Zanda County, Xizang Autonomous Region, China at 3300 m a.s.l.).
Remarks.The description of the species is based on the holotype, the only available voucher of the taxon (fig.8).

Diagnosis.
Comparatively large; head and body depressed; head scales mostly smooth; no enlarged, mucronate scales either on flanks or on body; higher number of scales around mid body; groups of low spinose scales on neck and sides of head; distinct tail segment of four whorls above becoming three below; callous glands both at precloacal and abdominal position.
Description.Head and body depressed; snout more than one and half times the distance between eye-tympanum or eye width and more than two times that of tympanum diameter; tympanum exposed, more than half of eye width; nostril pierced on canthus rostralis in the form of unique diagonal slit like aperture, more than half of nasal, pointing outward and backward, touching rostral; no gular pouch, gular plicate; upper head scales heterogeneous, subequal, mostly smooth; labials 9 -10 (10 ± 0.4); groups of low spinose scales present on the neck and sides of head especially around tympanum, preauricle in series otherwise in groups; nuchal dentition clearly visible; vertebral scales keeled, larger than other small dorsals, arranged in 14 -16 rows of keeled scales; complete absence of any enlarged scales on the lateral sides of the body or flanks; other small dorsals distinctly smaller than enlarged ones; ventral scales smooth, smaller than enlarged vertebral; gular scales granular, smooth; skin of the lateral sides does not form distinct dorsolateral fold; total number of scales around midbody 180 -196 side; 28 -46 (36 ± 4) scales in first complete whorl around the tail; callous glands present in males only at precloacal and at abdominal position, number of rows at precloacal position 4 -6 and an oblong patch on abdomen.
Head pale yellow or grey, sometimes speckled, on the sides black bar between eye and tympanum; gular spotted or speckled with black, males marked with blue, in juveniles reticulated; chest and belly pale yellow but may be speckled with age, males with blue or grey wash; body colour highly variable, sometimes dust brown or grey without any noticeable pattern, may be olive brown mottled with black and yellow, yellow spots may be of different sizes, the bigger ones are sometimes arranged in transverse series; another pattern which is generally observed in females is characterized as olive brown, light vertebral stripe with lateral extensions which are bordered by black and several other light and dark spots and specks, sub-adults have irregular pattern; tail shows indistinct banded pattern but when the specimen itself lacks any pattern on the body, the tail is also devoid of it.
Taxonomy.duda (1966, 1972, 1974) carried out morphological and anatomical studies, while BöhMe (1988) found strong differences in hemipenes morphology of L. tuberculata (the type species of Laudakia) as compared to Paralaudakia himalayana and P. lehmanni.These hemipenial differences support the distinct character of Laudakia and Paralaudakia.

Distribution. Eastern Afghanistan, northwestern
Pakistan, Kashmir, some parts of Uttar Pradesh, India, and southwestern Nepal.
Laudakia tuberculata has a very wide distribution.In 1827 it was originally described from Bengal, without specifying any precise locality.At that time Bengal was a very large province and as L. tuberculata is a mountain dwelling species, it is very likely that the type series was collected in northern parts of the province.BoulenGer (1885) synonymized Barycephalus sy ke sii and Stellio indicus with L. tuberculata, including several new localities (Tibet and Bengal (?) in the East, Simla, Kashmir, Ladakh, and Murree in the West).Laudakia tuberculata is widely distributed in Pakistan and the Kashmir area now associated with Pa kistan.Its presence in eastern Afghanistan is very likely because of a continous mountain range which spreads from Pakistan into Afghanistan.
The presence of both L. tuberculata and L. dayana is known from the Simla area of India, but it is recently unknown if both species are in true sympatry in this area.Because of altitudinal distributions in mountain areas, the presence of two taxa does not necessarily imply a sympatric occurence.It can be supposed that in Simla area L. tuberculata is distributed in lower Diagnosis.Closely related in morphology to Laudakia sacra but the latter taxon has the lateral scales uniform in size and not intermixed with large scales.
Description.Body depressed, head slightly triangular, longer than wide.Nasal elliptic, connected with the rostral and first supralabial, nostril in the center, directing outwards, nostril large, oval, just below the canthus rostralis, 4 to 5 small scales between posterior margin of nasal and superciliary ridge.Superciliary ridge not well developed.Eye moderate with round pupil, upper and lower eyelids covered with scales, altitudes while L. dayana occupy higher elevations.Specimens (e.g.ZMA 11636) identified as L. dayana were collected from the bank of Sutlaj River in upper Simla at an elevation of 3000 m a.s.l.while L. tu berculata is known from elevations between 1200 -2200 m in Pakistan and Kashmir.Waltner (1991) studied the altitudinal ecology of L. tuberculata in the Utter Pradesh state of India and found several dissimilarities among the populations living at different altitudes.However, it could be possible that one population belongs to L. tuberculata while others to L. dayana, but that has yet to be examined.
Waltner (1991) defined the eastern distribution limit up to Kathmandu (Nepal), but the records show the presence up to Sikkim.Few specimens of L. tubercu la ta have been reported from Singapur (BMNH xxiii.536) and Burma (ZMB 6052), but these are doubtful.

Fig. 3a .
Fig. 3a.Skulls of L. pakistanica, L. agrorensis and L. stellio.Left column= dorsal view of the skull; middle column= ventral view of the skull; right column= lower jaws from above and left.L. pakistanica and L. agrorensis are originally published in BaiG (1992), whereas L. stellio is obtained from el-touBi (1947a).
s. daani, the status of these populations remains open.The eastern boundary of the range remains open.Habitat.According to Franzen et al. (2008) exposed rocks, walls or screes from coastal to montane areas.

Distribution.
Northeastern Iran, mainly around Mashhad, some parts of Turkmenia and Afghanistan.In Turkmenistan the species is recognized from Badkhyz and Karabil in the south of the Tejen-Murgab rivers interstream area as well as in eastern Kopet Dagh.Its supposed subspecies (see taxonomy) was known only from eastern Kopetdag.ClarK (1991) reported P. erythrogaster from Torbat-i-Jam (el.1120 m) and an area about 60 km S.E. of Mashhad (1380 m) in Iran.Previously it was thought that it is restricted to northeastern Iran and southeastern Turkmenia at an elevation of 1000 -1700 m, but reports by anderson & leviton (1969), ClarK (1991) and BaiG (1992) indicate its presence also in Paktia and Paghman of eastern Afghanistan.The elevation records of anderson & leviton (1969) have expanded the vertical range of this species up to 2440 m.
strongly plicate; upper head scales heterogeneous, subequal, smooth above but spinose posteriorly; labials 10 -12 (11 ± 1.0); groups of spinose scales present on neck and sides of head especially around tympanum; vertebral scales heterogeneous, irregular, keeled, larger than other dorsal scales, midvertebral moderately large and partly interrupted by transverse rows of highly enlarged mucronate scales, sometimes giving impression of transverse folds on the body; no true patch of enlarged mucronate scales on flanks but groups of spinose scales randomly present over the sides of body; other small dorsals distinctly smaller than enlarged ones; ventral scales flat with posterior spiny tip, smaller than enlarged vertebral & flank scales; gular scales mucronate or spinose, those in the middle distinctly enlarged; total number of 1843 Stellio cyprius FitzinGer (nomen nudum fide anderson 1898), Syst.Rept.1: 85.Type locality: "Asia.Ins.Cyprus."1879 Stellio cordylina Günther (nomen nudum), Proc.Zool.Soc., London: 741.Type locality: "Cyprus."1967 Agama stellio cypriaca daan, Beaufortia 14(172): 127.Type locality: "Limasol, Cyprus."Diagnosis.Head only slightly depressed with swollen cheeks; vertebral scales heterogeneous, irregular, keeled, larger than other dorsal scales, mid-vertebral scales moderately large and sometimes interrupted by transverse rows of highly enlarged mucronate scales; tail at least one and half times the distance of snoutvent; yellowish vertebral bloches rarely visible; tail segment of two which changes into three in distal half; 3 -5 rows of precloacal callous glands and a stripe at the abdomen in males only.Description.Head heavy, only slightly depressed; snout longer than the distance between eye-tympanum or eye width; tympanum exposed, almost equal to eye width; nostril pierced below canthus rostralis, less than half of nasal scale in size, touching rostral; no true gular pouch but shows marked tendency as compared to all remaining taxa of 'Laudakia' in the old sense; gular scales around midbody 127 -168 (145 ± 9.1 [in examined juveniles number was less than 130]); limbs very strong, covered with enlarged mucronate scales, hindlimb slightly longer than the distance between gular fold and cloaca; fingers and toes compressed, 17 -21 (18.8 ± 1.3) lamellae under 3rd finger and 20 -27 (24.2 ± 1.6) under 4th toe.Tail almost rounded; each tail segment consists of two whorls of enlarged mucronate scales which change into three in the distal half; Larger size than other subspecies; head and body moderately depressed, head dorsally coloured like back or darker, not distinctly coloured from back, back with usually 4 -5 yellowish vertebral blotches on dark grey ground color, belly whitish, throat whitish ground colour, spotted dark usually over half of the area; vertebral scales heterogeneous, irregular, keeled, larger than other small dorsals, midvertebral moderately large and interrupted at regular intervals by transverse series of highly enlarged mucronate scales; 3 -5 rows of precloacal callous glands with more than 30 scales; tail segment of two whorls, tail length about one and half times the distance of snout-vent.
1980 Agama stellio daani Beutler & Frör, Mitt.Zool.Ges.Braunau 3: 270 -272.Holotype: ZSM 201/1978/2 (originally ZSM 201/ 1978 -1).Type locality: "Zw.Agh.Kirikos u.Evdilos, Ikaria, Region Samos, Griechenland [= between Agh.Kirikos and Evdilos, Ikaria, Samos Region, Greece.]." Diagnosis.Description.Head and body moderately depressed; snout longer than the distance between eye-tympanum or eye width and more than twice that of tympanum diameter; tympanum exposed, more than half of eye width; nostril pierced below canthus rostralis, less than half of nasal scale in size, touching rostral; no gular pouch, gular plicate; upper head scales heterogeneous, subequal, smooth or carinated; gular scales heterogenous, mucronate to spinose; groups of spinose scales present on the neck and sides of head especially around tympanum, in groups; narrow vertebral zone of enlarged scales in up to six rows, vertebral scales heterogeneous, irregular, keeled, larger than other dorsal scales, mid-vertebral moderately large and interrupted at regular intervals by transverse rows of enlarged mucronate scales, these rows are interrupted by 5 to 6 rows of granular scales; ventral scales smooth, smaller than vertebral and flank scales; limbs strong, covered with enlarged mucronate scales, hind-limb slightly longer than distance between gular fold and cloaca; fingers and toes compressed, lamellae 16 -23 (19.7) under fourth toe; tail moderately depressed, oval in cross section; each tail segment consists of two whorls of enlarged mucronate scales; callous glands present in males at precloacal and also at abdominal position, number of rows at precloacal position 3 -5.
smaller than enlarged vertebral & flank scales; gular scales very small, spinose; total number of scales around midbody184 -194 (190 ± 3.9); limbs strong, covered with enlarged mucronate scales, hindlimb slightly longer than the distance between gular fold and cloaca; fingers and toes compressed, 15 lamellae under 3rd fingerand 18 -21 (19.8 ± 1.3) under 4th toe.Tail very depressed, oval in cross section; tail segment distinct, each segment consists of two whorls of enlarged mucronate scales; 21 -27 (24 ± 3.0) scales in first complete whorl around the tail; 5 rows of callous glands present in males at precloacal and a narrow stripe at abdominal position.Males jet black above with traces of yellowish spots on some dorsal scales, on the toes and on the back; chin and throat black, other lower parts brownish grey.Females are also black with yellow spots on head and body, those on the body are arranged in rather irregular transverse rows; tail banded in both sexes, anterior one or two bands are broken; gular reticulate; belly yellow speckled with black.Distribution.Black Lava Desert of Jordan, Syria and Saudi Arabia.
. 1. Type locality: "Black Lava Desert of Transjordania (32° 10' N × 36° 40' E)."Diagnosis.Smaller size than other subspecies; head and body much depressed; colour black; vertebral scales heterogeneous, irregular, keeled, larger than other small dorsals, mid-vertebral moderately large and interrupted at regular intervals by transverse rows of enlarged mucronate scales; tail segment of two scale whorls , tail length about 20% more than the distance of snout-vent.Description.Head and body much depressed; snout longer than the distance eye-tympanum or eye width and about twice that of tympanum diameter; tympanum exposed, almost equal to eye width; nostril pierced on edge of canthus rostralis, less than half of nasal scale in size, directed outward and backward; no gular pouch, gular plicate; upper head scales heterogeneous, subequal, mostly carinated; upper labials 10 -15 (12 ± 2.1) and lower 12 -13 (13 ± 1.0); groups of spinose scales present on the neck and sides of head especially around tympanum, in groups; narrow vertebral zone of enlarged scales, vertebral scales heterogeneous, irregular, keeled, larger than other small dorsals, mid-vertebral moderately large and interrupted at regular intervals by transverse rows of highly enlarged mucronate scales, transverse rows themselfs may not be regular; no true patch of enlarged mucronate scales on flanks but these scales are randomly present over the sides of body, sometimes in groups; other small dorsals distinctly smaller than enlarged ones; ventral scales smooth, 1951 Agama stellio brachydactyla haas (part), Ann.Mag.Nat.Hist., London, 12 (4): 1052.2006 Laudakia stellio salehi Werner, in: laChMann et al., J. Nat Hist.40: 1259 -1284.Type locality: "Sinai: 3 km from Sheikh Harun towards Watiya Pass, 28° 35' 30'' N, 33° 59' E, alt.1650m."

1896
Stellio erythrogaster niKolsKy, Annuire Mus.zool.Acad.Sci.St.Peterbourg, 1 App.: 370.Type local-Contrary to other Paralaudakia it lives in clay and sandy-loamy soils, mainly in areas with colonies of Rhombomys WaGner, 1841 (Rodentia: Muridae), and it avoids vertical slopes (terent'ev & Chernov 1949).ClarK et al. (1966) found them on man made rock piles in Iran, but ClarK (1991) characterized the habitat as deep holes and crevices in earth banks, cliffs, and piles of stones.Remarks.terent'ev&Chernov (1949) and ana nJe va et al. (1998) mentioned that P. erythrogaster uses rodent burrows as hideouts and feeds on insects, mainly beetles and caterpillars, and plants.anderson&leviton (1969) examined stomach contents of this species and found no plant material.foldandcloaca; fingers and toes compressed 18 -19 lamellae under third finger and 23 -27 under fourth toe, fourth toe considerably longer than third, extremity of the claw of the latter does not reach the base of the claw of former.Tail depressed, oval in cross section; each tail segment consists of three complete whorls of enlarged (largest of all body scales), spinose scales; 26 -32 scales in first complete whorl around the tail; callous glands present in both sexes, in males 3 -5 rows at precloacal and a large patch at mid abdominal position, in females up to two rows at precloacal and rarely possess a small patch at abdomen.Head olive or grey with black spots over it; gular grey with or without yellow specks or reticulation; chest and belly pale yellow, in males with grey wash; vertebral stripe usually grey; olive grey above with dark rimmed yellow ocelli, males usually do not show bright and marked pattern; tail with dark spots which sometimes gives the impression of cross bars.observed that A. chernovi and A. iso zona both differ from P. himalayana, but they hardly show any dissimilarity with one another.ananJevaetal.(1981)distinguished A. chernovi from P. himala yana and P. badakshana by having a relatively longer tail, always keeled enlarged vertebrals, fewer number of enlarged vertebral rows, and more rows of callous precloacal scales.However, all these have been proved true with respect to A. isozona.Type material of P. bo chariensis was not examined but the description given by niKolsKy (1915) shows several common characteristics similar to A. isozona and A. chernovi.Moreover, all three (P.bochariensis, A. isozona and A. chernovi) had been described from areas in close proximity to one another.It is therefore, suggested that A. chernovi and A. isozona should be considered as a junior synonyms of P. bochariensis.

.
(ananJeva et al. 1998)and ananJeva et al.(1998): olive-grey, clayish or brown with vermiculate short black small spots and line bars.The northern border of the range passes along the foothills of the mountain range Mogoltau in the Fergana valley, the western border follows the mountain ranges Nuratau and Kugitang-Tau and is limited to the Darvaz range(ananJeva et al. 1998(ananJeva et al.  ,  2006)).anderson&leviton(1969)did not find any nearest record of P. himalayana is Sarhad (Wakhan, Afghanistan), south of Pamir.The locality of Schador close to Wakhan makes the presence of P. himalayana observed by ananJeva et al.(1981)further west or northwest of Pamir even more doubtful.Except for the forementioned records, all other records of P. himalay ana, P. badakshana and P. bochariensis (together with A. isozona and A. chernovi) are allopatric.However, populations of P. himalayana in Chitral (Pakistan) represent an intermediary position and show resemblance with P. badakshana in some characteristics (e.g.number of scales around the body).We assume that the records of P. himalayana from Afghanistan (anderson & leviton 1969) might belong to a forementioned Chitral population.Presence of P. himalayana in an unidentified area in Schador, observations regarding the sympatry of P.himalayana and A. chernovi (ananJeva et  al. 1981), and an intermediary population of Chitral do not allow us to consider these three taxa (P.himalaya na, P. badakshana and P. bochariensis) as subspecies.Therefore, we recognize them in a superspecies complex as follows: Paralaudakia (himalayana) hima layana, Paralaudakia (himalayana) bochariensis, and Paralaudakia (himalayana) badakshana.
Mertens (1969)lio liratus from Baluchistan and later BoulenGer (1885) distinguished it from L. melanura as having partly keeled head scales, six or seven rows of keeled vertebral scales and by the absence of caudal rings.sMith(1935)synonymized it with L. melanura and mentioned specimens with smooth enlarged vertebral scales from the northern range while specimens from the southern part have them more or less keeled.This might be partly true with respect to some ranges, but BaiG (1992) examined specimens with keeled enlarged vertebrals (but with very low ridges) and transversely keeled head scales near the nuchal end of head also in the northern part of the range (Salt Range, Pakistan), and smooth vertebral scales in specimens from southern parts of Sind and Baluchistan.Therefore, this is not a consistent character within the species and either habitat or age might have some influence on it.Mertens (1969)again resurrected L. lirata as a subspecies of L. melanura on the basis of carinated scales and colouration.However, both of these characters are apparently inconsistent and therefore should not be considered as sufficient evidence to split both taxa on subspecies level.
This form is very close to the nominate form in certain characteristics but quite distinguished by being dull dark brown to grey in colour with white transverse bands across the body and having a fewer number of scales around the body.
Remarks.This is an extremely wary species, which retreats quickly into a crevice when alarmed.Observations (particularly of those living near build- covered with strongly mucronate above and flat scales below, hindlimb longer than distance between gular fold and cloaca; 22 -26 lamellae under third finger and 26 -31 under fourth toe.Tail depressed, oval in cross section, forming very distinct annuli, each tail segment consists of three complete whorls; 33 -40 scales in first complete whorl around the tail, scales on dorsal and lateral sides strongly mucronate and spinose, spine extends slightly beyond the posterior margin of scale; ventral scales smooth. Laudakia tuberculata (HarDWicke & Gray, 1827) 1827 Agama tuberculata hardWiCKe & Gray, Zool.J. London, 3: 218.Type locality: "India," interpreted as "Bengal" by sMith 1935.1853StellioindicusBlyth (fide sMith 1935; non Agama indica Gray = Calotes versicolor), J. Asiat.Soc.Bengal, Calcutta, 22: 647.Type locality: "Mirzapore [= Wazirabad, upper Hindustan]." 1860 Barycephalus sykesii Günther (fide sMith 1935), Proc.zool.Soc.London, 1860: 150; pl. 25, fig. A. Type locality: "Simla, Himalaya, 2500 feet above sea level; Simla, Himalaya, 7200 feet above sea level; Gårhvál, Himalaya, 8200 feet above sea level; Balti, Tibet, 6100 feet above sea level; Ladak, Tibet, 15,250 feet above sea level," interpreted as "Simla, Himalaya" by sMith 1935.Head and body depressed; head scales smooth or rugose; no patch of enlarged, mucronate scales on flanks; only scattered, enlarged, mucronate scales present on body; groups of low spinose scales on neck and sides of head; larger number of scales around body; tail segment of four whorls which remain of three below, indistinct proximally; callous glands in males only.Description.Head and body depressed; snout one and half times or more the distance between eye-tympanum or eye width and more than twice that of tympanum diameter; tympanum exposed, more than half of eye width; nostril pierced on or below canthus rostralis, less than half of nasal, pointing outward and backward, touching rostral; no gular pouch, gular plicate; upper head scales heterogeneous, subequal, smooth or posteriorly slightly rough; labials 8 -12 (10 ± 1); groups of low spinose scales present on the neck and sides of head especially around tympanum, preauricle in series otherwise in groups; vertebral scales keeled, larger than other small dorsals, 12 -14 oblique rows of keeled scales; no patch of enlarged mucronate scales on flanks, enlarged mucronate scales scattered over the sides of body; other dorsals smaller than enlarged ones; ventral scales smooth, smaller than enlarged vertebral but larger than gular and other small dorsal scales; gular scales granular, smooth; skin of lateral sides loose forming dorsolateral fold; total number of scales around midbody134 -221 (171 ± 22.3); limbs strong, covered with enlarged mucronate scales, hind-imb slightly longer than distance between gular fold and cloaca; fingers and toes compressed, 20 -30 (24.8 ± 2.49) lamellae under third finger and 26 -37 (31.5 ± 2.6) under fourth toe.Tail depressed, oval in cross section; each tail segment consists of 4 whorls of enlarged mucronate scales but remain 3 on ventral Laudakia wui zHao, 1998 1998 Laudakia wui zhao, Acta Zootax.Sin.23: 440 -444.Type locality: "Yi'ong, Bomi County, Xizang [Tibet] Autonomous Region, China; altitude 2350 m." Diagnosis.