RESEARCH ON ASIAN AGAMID LIZARDS: CONTRIBUTION OF ZOOLOGICAL INSTITUTE, RUSSIAN ACADEMY OF SCIENCES, ST. PETERSBURG

Russian Journal of Herpetology Vol. 19, No. 2, 2012, pp. 85 100 RESEARCH ON ASIAN AGAMID LIZARDS: CONTRIBUTION OF ZOOLOGICAL INSTITUTE, RUSSIAN ACADEMY OF SCIENCES, ST. PETERSBURG Natalia B. Ananjeva 1 Submitted May 14, 2011. There is an overview of the study of agamid lizards in the Zoological Institute of the Russian Academy of Sciences (ZISP), St. Petersburg, which is inseparably associated with general zoological research in Russia. During the period of exploration and organized scientific expeditions, the Russian Academy of Sciences aimed to study the natural resources of the unbounded territories of Russia, and a number of prominent foreign scientists were invited to Russia for these research expeditions. One of the most prominent scientists was Peter S. Pallas (1741 1811), whose activities resulted in the first inventory of biological resources of the country. Contributions of A. Strauch who was a founder of the Russian school of herpetology, J. von Bedriaga. A. M. Nikolsky, S. A. Chernov, I. S. Darevsky and other leading Russian and Soviet herpetologists are considered. I also try to provide the overview of the current research of Asian agamids in Zoological Institute of the Russian Academy of Sciences. Keywords: Squamata; Acrodonta; Agamidae; history; Department of Herpetology, Zoological Institute, St. Petersburg. INTRODUCTION Pallas and earlier explorations The first species in the genus Phrynocephalus discovered by zoologists were referred to the catch-all lizard genera Lacerta (Pallas, 1771, 1776; Lepechin, 1774; Gmelin, 1789; Pallas, [1814]) and Agama (Olivier, 1807; Lichtenstein in Eversmann, 1823) (Barabanov and Ananjeva, 2007). Among agamid lizards described by P. S. Pallas there are such species as Lacerta helioscopa Pallas, 1771: 457; Lacerta mystacea Pallas, 1776: 702; Lacerta aurita Pallas, 1776: Tab. V, Fig. 1; Lacerta caudivolvula Pallas [1814]: 27; Trapelus sanguinolentus [(Pallas 1814)] (see information about type specimens in Denzer et al., 1997; Barabanov and Ananjeva, 2007; Barabanov and Milto, 2012). The latter species, steppe agama, was described from the sands of Kum-Ankatar in the valley of the Terek River, referred to as Lacerta agama Gyuldenshtedt (Guldenstaedt) in his field diaries (Ananjeva and Tsaruk, 1987). Later, P. S. Pallas included the previously unpublished description in Volume III of Zoographia Rosso- Asiatica [1814], giving to a new lizard a new species name, Lacerta sanguinolenta, instead of the author s 1 Zoological Institute, Russian Academy of Sciences, Universitetskaya nab., 1, 199034 St. Petersburg, Russia; e-mail: azemiops@zin. ru name, preoccupied by Lacerta agama L., 1758 (currently Agama agama L., 1758). In the text, there is a direct link to the authorship of Johann Anton Güldenstädt (p. 23): «¹ 19 Lacerta sanguinolenta tab. IV, Fig. 2. Lacerta agama Güldenst. not Ms. Species insigmen a me non obscuratam in collibus arenosis Kum-Ankatar dictis ad Terek fl. invenit Guldenstaedt et discriptionem cum icone reliquit». Authorship of Güldenstädt is also confirmed by Lichtenstein (Lichtenstein in Eversmann, 1823: 145). Famous naturalists, collectors and explorers associated with the Zoological Institute such as S. G. Gmelin and E. Eichwald, have made great contributions to the study of the herpetofauna of the Russian Empire. A number of agamid taxa names were proposed by Gmelin: Lacerta uralensis Gmelin, 1789: 1073, Lacerta guttata Gmelin, 1789: 1078, by Lichtenstein: Agama ocellata Lichtenstein in Eversmann, 1823: 143 and by Eichwald: Phrynocephalus varius Eichwald, 1831: 186, Phrynocephalus reticulatus Eichwald, 1831: 186, Phrynocephalus melanurus Eichwald, 1831: 186, and Phrynocephalus nigricans Eichwald, 1831: 186. Eichwald (1831) also proposed the new generic name Megalochilus for Phrynocephalus mystaceus (Fig. 1) and described from the Caucasus the wide spread Eurasian species Stellio caucasius Eichwald, 1831. 1026-2296 2012 1902-0085 2012 Folium Publishing Company

86 Natalia B. Ananjeva Fig. 1. Drawings of Megalochilus auritus and Trapelus sanguinolentus from Eichwald (1841). At first the Caucasian agama was described under the name Lacerta muricata by Pallas in 1814. This name however, could not be used as valid because earlier (Shaw, 1802), another agamid lizard (known now as Amphibolurus muricatus) was described under the same name. To identify the form described by P. S. Pallas as the Caucasian agama is now possible only from the description itself, or by references of that time, as unfortunately, no specimens of lizard or its drawing remain. No image of Lacerta muricata, as it is generally referred to, was published in the color tables of the drawings to Zoographia Rosso-Asiatica. It is also not registered in collections of the Archive of the Russian Academy of Sciences, which are stored now as a part of the materials by P. S. Pallas. In 1831, E. Eichwald described the Caucasian agama under the name Stellio caucasius, which was used in the herpetological literature. In 1841 Eichwald returned to this lizard in his fundamental work «Fauna Caspio-Caucasia» (1841) (Fig. 2) and gave a first detailed description of the external morphology and osteology, as well as the image of the skeleton, the skull alone, the lower jaw, and pectoral girdle (Table III, Fig. 108) (Fig. 3). From this book (Fig. 4) and Eversmann (1834) Fig. 2. Eichwald E. (1841), Fauna Caspio-Caucasia nonnullis observationibus novis, Petropoli. Cover page. (Fig. 5) we know the drawing of Phrynocephalus caudivolvulus (= P. guttatus) (Fig. 6). Eversmann (1834) provided also early drawings of P. helioscopus and Stellio vulgaris (Fig. 6). He noted this latter species in the southern regions of the Caucasus; in fact it should be Laudakia caucasia. Agamid Studies in 19 th Century Alexander A. Strauch, academician and museum director (from 1879 to 1890) was a founder of the Russian school of herpetology. As a result of his own careful study of turtles, crocodiles, lizards, snakes, and salamanders, Strauch wrote several monographs and world synopses (Strauch, 1866, 1869, 1870, 1873, 1887, 1890). Among his publications is the first catalogue of the herpetological collection in the Zoological Museum, which included 5889 catalogue numbers representing 1222 species of reptiles (119 species of turtles, 14 crocodiles, 596 lizards, and 493 snakes) and 1285 catalogue numbers of 283 species of amphibians (9 caecilians, 224 frogs, and 50 salamanders). This collection was widely representative for its time (Strauch, 1889). The results of

Research on Asian Agamid Lizards: Contribution of Zoological Institute, Russian Academy of Sciences 87 Fig. 3. Skeleton of Stellio caucasia (1 8) and Phrynocephalus caudivolvulus (9 14) from Eichwald (1841). Fig. 5. Eversmann E. (1834), Lacertae Imperii Rossici variis in itineribus meis observatae, Nouv. Mem. Soc. Natur. Moscou. Cover page. Fig. 4. Drawings of Phrynocephalus caudivolvulus and Ophisops elegans from Eichwald (1841). Strauch s identification and description of the amphibians and reptiles (1876) from Przewalski s first expedition to Central Asia are of special value for Russian and world herpetology. Strauch also proposed the following names of taxa of agamid lizards: Centrotrachelus asmussii Strauch 1863: 479) (Fig. 7), Phrynocephalus affinis Strauch 1876: 13), Phrynocephalus frontalis Strauch, 1876: 15, Phrynocephalus przewalskii Strauch (1876: 10), Phrynocephalus vlangalii Strauch, 1876: 20, Phrynocephalus versicolor Strauch, 1876: 18, and Phrynocephalus frontalis Strauch (1876: 15). A very significant source of new and interesting materials on arid Asian agamids originated from the first expedition of the famous explorer Nikolai M. Przewalski in the vast unknown and remote regions of Central Asia, along with other arid reptiles. Discoveries of toad-headed agamas of the genus Phrynocephalus were of special importance. The well-known herpetologist, one of founders of Russian herpetology and researchers of the fauna of deserts of Central Asia, Jacques von Bedriaga who was Fig. 6. Drawings of Stellio vulgaris (1), Phrynocephalus caudivolvulus (2), and P. helioscopus (3) from Eversmann (1834). informally associated with the Zoological Museum in St. Petersburg (Adler, 1989; Ananjeva, 1998, 2005), made his famous and important contribution as a large monograph (in quarto and nearly 800 pages) (Bedriaga,

88 Natalia B. Ananjeva Fig. 7. Type specimen ZISP 3029. Centrotrachelus asmussii Strauch, 1863. 1898, 1905 1912) devoted to the study of the huge herpetological collections made by Przewalski and other Russian explorers during their Central Asian expeditions (Figs. 8 19). He called this taxonomically complicated group of Palaearctic lizards the most difficult task with which they can deal herpetologists. His generic system, proposed in 1907, which allowed dividing the genus into large groups, was based on an evaluation of the relative length of internasal distance. On the basis of the richest collections of Central Asian agamids, Bedriaga suggested 42 names for agamids, all of them for Phrynocephalus: Phrynocephalus acutirostris Bedriaga 1909: 339 Phrynocephalus alpherakii Bedriaga in Nikolsky 1905: 470 Phrynocephalus arcellazzii Bedriaga 1909: 307 Phrynocephalus axillaris var. ericae Bedriaga 1905 1907a: 168 Fig. 8. Bedriaga J. (1898), Amphibien und Reptilien (Reptilia Przewalskiana), in: Wissenschaftliche Resultate der von N. M. Przewalski nach Central-Asien unternommenen Reisen. Zoologischer Theil, Bd. 3, Abth. 1. Cover page. Phrynocephalus axillaris var. idae Bedriaga 1905 1907a: 159 Phrynocephalus axillaris var. klemenzi Bedriaga 1907: 163 Phrynocephalus axillaris var. maculata Bedriaga 1907: figure caption to the Table III, Fig. 3 Phrynocephalus axillaris var. murielis Bedriaga 1905 1907a: 170 Phrynocephalus blanfordi Bedriaga 1909: 381 Phrynocephalus boulengeri Bedriaga in Nikolsky 1905: 466 Phrynocephalus carinilabris Bedriaga 1909: 412 Phrynocephalus caudivolvulus var. incerta Bedriaga in Nikolsky 1905: 475 Phrynocephalus ciliaris Bedriaga 1909: 297 Phrynocephalus grum-grzimailoi Bedriaga 1909: 420 Phrynocephalus guentheri Bedriaga 1905 1907a: 168 Phrynocephalus haeckeli Bedriaga 1907: 237

Research on Asian Agamid Lizards: Contribution of Zoological Institute, Russian Academy of Sciences 89 Fig. 10. Captions to drawings in Table II from Bedriaga (1898). Fig. 9. Table II from Bedriaga (1898). Phrynocephalus helioscopus var. cameranoi Bedriaga 1907: 243 Phrynocephalus helioscopus var. orientalis Bedriaga 1912: 751 Phrynocephalus isseli Bedriaga 1907: 229 Phrynocephalus koslowi Bedriaga 1905 1907a: 162 Phrynocephalus kuschakewitschi Bedriaga in Nikolsky 1905: 476 Phrynocephalus nikolskii Bedriaga in Nikolsky 1905: 463 Phrynocephalus pewzowi Bedriaga 1907: 139, 173 Phrynocephalus potanini Bedriaga 1909: 389 Phrynocephalus putjatai Bedriaga 1909: 302 Phrynocephalus raddei var. boettgeri Bedriaga in Nikolsky 1905: 468 Phrynocephalus reldoe Bedriaga 1909: 495 Phrynocephalus salenskyi Bedriaga 1907: 141, 213 Phrynocephalus steindachneri Bedriaga 1907: 140, 207 Phrynocephalus suschkini Bedriaga in Nikolsky 1905: 472 Phrynocephalus versicolor var. bogdanowi Bedriaga 1909: 334 Fig. 11. Table IV from Bedriaga (1898).

90 Natalia B. Ananjeva Fig. 12. Captions to drawings in Table IV from Bedriaga (1898). Fig. 14. Bedriaga J. (1909), Amphibien und Reptilien (Reptilia Przewalskiana), in: Wissenschaftliche Resultate der von N. M. Przewalski nach Central-Asien unternommenen Reisen. Zoologischer Theil, Bd. 3, Abth. 3. Cover page. Fig. 13. Captions to drawings in Table IV from Bedriaga (1898). Phrynocephalus versicolor var. doriai Bedriaga 1909: 332 Phrynocephalus versicolor var. hispida Bedriaga 1909: 331 Phrynocephalus versicolor var. kulagini Bedriaga 1909: 329 Phrynocephalus versicolor var. siebenrocki Bedriaga 1909: 333 Phrynocephalus vlangalii var. geckoides Bedriaga 1909: 466 Phrynocephalus vlangalii var. grombtschewskii Bedriaga 1909: 463 Phrynocephalus vlangalii var. lidskii Bedriaga 1909: 479 Phrynocephalus vlangalii var. nanschanica Bedriaga 1905 1907a: 172 Phrynocephalus vlangalii var. parva Bedriaga 1909: 468 Phrynocephalus vlangalii var. pylzowi Bedriaga 1909: 471 Phrynocephalus vlangalii var. roborowskii Bedriaga 1905 1907a: 172 (Fig. 20)

Research on Asian Agamid Lizards: Contribution of Zoological Institute, Russian Academy of Sciences 91 Fig. 16. Captions to drawings in Table VII from Bedriaga (1909). Fig. 15. Table VII from Bedriaga (1909). The next stage of development of herpetology in St. Petersburg was closely related to activities of the leading Russian and Soviet herpetologist, Alexander M. Nikolsky (1858 1942) who was a successor to Alexander Strauch as custodian of the herpetological collection at the Zoological Institute. Nikolsky was the author of several important monographs with information of continued importance, including synopses of the herpetofauna of Turkestan (1899) and the Caucasus (1913), and the first comprehensive books on the amphibians and reptiles «Fauna of Russia and Adjacent Countries» in 3 volumes, 1915, 1916, 1918) (Figs. 21 23). Another important contribution of Nikolsky was the study, identification, and description of the rich herpetological material collected in Iran by Nikolai Zarudny who was one the first zoologists to study the diverse fauna of Persia (Anderson, 1999; Ananjeva, 2008). In 1896 1904 N. A. Zarudny conducted four expeditions, famous worldwide, to Persia (1896, 1898, 1900 1901, 1903 1904) supported by the Russian Academy of Sciences, the Russian Geographic Society and St. Petersburg Zoological Museum. Nikolsky described Agama reticulata Nikolskij 1911: 272, Stellio erythrogaster Nikolskij, 1896: 370, Stellio erythrogaster var. pallida Nikolskij, 1897: 319, Stellio lehmanni Nikolskij, 1896a: App. XIV, Stellio bochariensis Nikolskij, 1897: 159: Agama kirmanensis Nikolskij, 1899 1900: 389, Agama kirmanensis var. brevicauda Nikolskij, 1907: 272, Phrynocephalus spiniventris Nikolskij, 1896: 370, Phrynocephalus strauchi Nikolskij, 1899: 21, Phrynocephalus rossikowi Nikolskij, 1899a: 286, Phrynocephalus olivieri var. brevipes Nikolsky, 1907: 274, Phrynocephalus olivieri var. carinipes Nikolskij, 1907: 273, Phrynocephalus caudivolvulus var. moltschanovi Nikolskij, 1913: 34, Phrynocephalus raddei var. zarudnyi Nikolskij, 1915: 193, Phrynocephalus rossikowi relictus Nikolskij 1915: 169, Phrynocephalus raddei var. bilkewitschi Nikolskij, 1915: 194 (Wermuth, 1967; Barabanov and Ananjeva, 2007; Milto and Barabanov, 2012). Interesting to note that Nikolsky and Zarudny carried out the joint trip to the Transcaspian region and north-eastern Persia in 1885. Further development of the taxonomy and systematics of the toad-headed agamas is associated with the activity of another well-known herpetologist and curator of herpetology, Sergey F. Tzarewsky (Garanin, 2009; Adler, 2012; Ananjeva and Doronin, 2012). After organization of the herpetological department as an administrative unit, it was headed by Tzarewsky (1919 1929), who

92 Natalia B. Ananjeva Fig. 17. Bedriaga J. (1912), Amphibien und Reptilien (Reptilia Przewalskiana), in: Wissenschaftliche Resultate der von N. M. Przewalski nach Central-Asien unternommenen Reisen. Zoologischer Theil, Bd. 3, Abth. 4. Cover page. Fig. 18. Table IX from Bedriaga (1912). published, in particular, articles on the taxonomy of the boas (Eryx) and toad-headed agamas (Phrynocephalus) which are of serious value. Tzarewsky worked at the Zoological Museum of the Academy of Sciences until 1930, when he was arrested. After his arrest in 1930 and subsequent release in five years, he left Leningrad and was not further involved in zoological activity. In 1964, at the request of P. V. Terentjev and V. I. Garanin, he published his last work in the abstract book of the First Herpetological Conference in Leningrad (Tzarevsky, 1964). Tzarewsky suggested the following names of toad-headed agamids (Barabanov and Ananjeva, 2007): Phrynocephalus bedrjagai Tzarewsky, 1926: 214. Phrynocephalus birulai Tzarewsky, 1927b: 304 Phrynocephalus suschkinianus Tzarewsky, 1927b: 304. Phrynocephalus carinatus Tzarewsky, 1927b: 305. Phrynocephalus elegans Tzarewsky, 1927b: 305. Phrynocephalus parvulus Tzarewsky, 1927b: 305. Phrynocephalus rostralis Tzarewsky, 1930: 216. Fig. 19. Captions to drawings in Table IX from from Bedriaga (1912). Tsarevsky focused his study on the structure of the skulls (Tsarevsky, 1926, 1927a, 1927b, 1929, 1964 and 1929 facsimile publication, this volume), suggesting that the system of diagnostic features proposed by

Research on Asian Agamid Lizards: Contribution of Zoological Institute, Russian Academy of Sciences 93 Fig. 20. Type specimen ZISP 7060. Phrynocephalus vlangalii var. roborowskii Bedriaga, 1907. Fig. 21. Nikolsky A. M. (1915), The Fauna of Russia and Adjacent Countries. Reptilia. Vol. I. Chelonia and Sauria, Academy of Science, Petrograd [in Russian]. Cover page. Bedriaga and based on the ratio of the width between the nasal openings and the distance from the front edge of the eye to the nasal aperture is insufficient for complex diagnostics (Fig. 24). Unfortunately, in general, these were species that are diagnostically well distinguished by other morphological characters, and he failed to establish a system of craniological characters for identification of poorly distinguished species. Soviet Period Successors of these great zoologists in the position of Head and Curator of the Department of Herpetology at the Zoological Institute, USSR Academy of Sciences were Sergei A. Chernov (1930 1960) and Ilya S. Darevsky (1961 1995). Both of them made a significant contribution to the study of agamid lizards. Sergei A. Chernov (1903 1964), a student of A. M. Nikolsky, was born in Kharkov, in the Ukraine and graduated from the University of Kharkov in 1926. In Kharkov he had been a student of A. M. Nikolsky. Chernov was appointed curator of the Department of Herpetology of the Zoological Institute at the Academy of Sciences of USSR, succeeding Sergei F. Tzarewsky. At that time the museum s name was changed to the Zoological Institute in 1930. Chernov, who become one of the leading Soviet herpetologists during his 30-year curatorship (1930 1960) conducted extensive herpetological field work in the Transcaspian region (1932), the Caucasus (1937 1939) and in Tajikistan (1942 1944). The results were published as a series of articles and several monographs, including regional synopses of the herpetofauna of Armenia (1937, 1939) and Tajikistan (1959). His most widely-known work was a book Field Guide to the Reptiles and Amphibians of the USSR, co-authored with Pavel V. Terentyev (three editions in 1936, 1940, 1949) that was translated into English in 1965. Chernov made a significant contribution to the study of the agamids he described Phrynocephalus interscapularis sogdianus Chernov (1948: 135) and Agama paw-

94 Natalia B. Ananjeva Fig. 22. Drawing of Phrynocephalus mystaceus from Nikolsky (1915). Fig. 23. Drawings of agamids and geckos from Nikolsky (1915). lowskii Chernov et Dubinin (1946: 741, 747) (see facsimile publication, this volume). The latter type the enigmatic agama (Fig. 25), described and known only from a single specimen. Several expeditions were organized to find more specimens however without any success. This species was later synonymized with African agama Agama mossambica (Ananjeva and Peters, 1981). It should be noted that the author of the description emphasized the differences of his species from all known Palaearctic agamas, which can indirectly confirm the African origin of this mysterious discovery. Occurrence of this species in the Middle Asia was never confirmed by repeated records in the same location (Fig. 26), in Khavast area on the border of Uzbekistan and Tajikistan (Ananjeva and Peters, 1981). Besides descriptions of new species, Chernov, in his monograph on the herpetological fauna of Tajikistan, was making a new attempt at taxonomic revision of the genus Phrynocephalus and published comparative remarks on the taxonomic status of the complex «Agama agilis» [A. (Trapelus) agilis, A. (Trapelus) sanguinolentus, A. (Trapelus) isolepis, A. (Trapelus) kirmanensis], which confirmed the species status of A. (Trapelus) sanguinolentus. The forms A. (Trapelus) isolepis and A. (Trapelus) kirmanensis were synonymized with A. (Trapelus) sanguinolentus. It should be noted that DNA analysis recently confirmed the correctness of S. A. Chernov opinion (Macey et al., 2000; Ananjeva, 2011). Critically analyzing the characters of toad-headed agamids, which were used as diagnostic characters by Bedriaga, Nikolsky and Tzarevsky, Chernov concluded that most of them are not specific. He gives the fundamental taxonomic value to the structure of toes, subdigital plates and the adjacent scales on the surface of lateral fingers. He developed identification keys to the species of Phrynocephalus, known from Middle Asia based on these characters. Chernov developed this system of diagnostic characters later in the identification keys in Field Guide of the amphibians and reptiles of the USSR (Terentyev and Chernov, 1949).

Research on Asian Agamid Lizards: Contribution of Zoological Institute, Russian Academy of Sciences 95 Fig. 24. Paper of Tzarevsky (Carevskij) S. F. (1927a), Contributions à la systématique et la distribution des lézards du genre Phrynocephalus (Reptilia), III. Comptes Rendus l Acad. Sci. l URSS, 23 26 [in Russian with French title]: on the left, hand-written manuscript; on the right, published paper. In 1962, Ilya Darevsky, a student of S. Chernov, became a third curator of herpetology (after Tzarewsky and Chernov) at the Zoological Institute of the USSR Academy of Sciences; in 1967, he brilliantly defended his Doctoral thesis (second scientific degree in Russia) The Rock Lizards of the Caucasus and in 1976, he became the Head of the Laboratory of Ornithology and Herpetology, which he led for the next twenty years. Later on, studies of the Caucasian herpetofauna, and the rock lizards in particular, made him a world-renowned scientist. He was the first to discover parthenogenesis and polyploidy in higher vertebrates and showed their importance for speciation. These discoveries inspired an explosion of similar investigations across many taxa and made Darevsky a world authority on the evolutionary importance of parthenogenesis and polyploidy in vertebrates. Among his numerous herpetological contributions is a description of the agamid Phrynocephalus reticulatus bannikovi Darevsky, Rustamov et Shammakov, 1976: 114. Working in the Caucasus, Darevsky (1960) made one of the most successful ecological studies of Phrynocephalus, on their population dynamics, movement and growth in the Arax River valley in Armenia. In 1957 he described specialized glands in the dorsal skin of this species. Subsequently, the glandular nature of these structures has not been confirmed: they were identified as skin tubercles (Sokolov et al., 1997), consisting of massive bases loaded in the cup-shaped epidermal depression, in the bottom of which are dermal papilla, and sensilla, prominent above the surface of the epidermis. They perform not only a receptor function (Ananjeva et al., 1991), but also serve as a protective, damping devices. Later, he was one of the first herpetologists in the Soviet-Vietnamese cooperation program, launched in 1978 to study the biota of tropical forests in Vietnam. As the author of the discovery of parthenogenesis in reptiles in the Caucasian rock lizards, he continued to develop this field to study tropical lizards, geckos and agamids. In co-authorship with L. A. Kupriyanova they described two parthenogenetic species: Leiolepis guentherpetersi Darevsky et Kupriyanova, 1993. Herpetozoa, 6(1/2), 9 Leiolepis boehmei Darevsky et Kupriyanova, 1993. Herpetozoa, 6(1/2), 4 We could not provide the most complete panorama of the study of the agamas without referring to our colleague from the Museum fur Naturkunde an der Humboldt-Universität zu Berlin, DDR, Günther Peters. This author of taxonomic revision of Central Asian toadheaded agamids of the genus Phrynocephalus, examined the extensive collections of the Zoological Institute, including important type specimens, and proposed his classification (Peters, 1984). In the course of this work he described the following species: Phrynocephalus steynegeri Peters, 1984: 37 Phrynocephalus werneri Peters, 1984: 39.

96 Natalia B. Ananjeva Fig. 25. Drawing of Agama pawlowskii Chernov et Dubinin, 1946 (artist Sergey M. Steinberg). Phrynocephalus vlangalii var. occidentalis Peters, 1984: 47. Phr[ynocephalus]. alticola Peters, 1984: 58. Moreover, in co-authorship with Natalia Ananjeva, G. Peters described a new species from Tajikistan, Agama chernovi Ananjeva and Peters et Rzepakovsky (1981: 23), which was later synonymized with Laudakia bochariensis (Nikolsky, 1897), and published taxonomic papers on the status of Agama pawlowskii and Laudakia sacra. Swen Veldre (1964) studied geographic variation of the widespread species. Phrynocephalus mystaceus, and these results confirmed the opinion of Pavel V. Terentyev, who rejected the existence of subspecies. Current Research Beginning in 1996, the curatorship of the Department of Herpetology and the administration of the Laboratory of Herpetology and Ornithology passed to Natalia B. Ananjeva (Ananjeva, 1998, 2005; Ananjeva and Darevsky, 2004), who holds these positions currently. She studied and defenses doctoral dissertation on the Palearctic agamids (1992) and later took part in the study on biodiversity of Vietnam together with N. L. Orlov and Vietnamese herpetologists (Nguyen, 2006). They described the following species of draconine agamids of Vietnam: Acanthosaura nataliae Orlov, Nguyen et Nguyen, 2006. Russ. J. Herpetol., 13(1), 62. Bronchocela vietnamensis Hallermann et Orlov, 2005. Russ. J. Herpetol., 12(3), 176. Pseudocophotis kontumensis Ananjeva, Orlov, Nguyen et Nazarov, 2007. Russ. J. Herpetol., 14(2), 154. Pseudocalotes ziegleri Hallermann, Nguyen, Orlov et Ananjeva, 2010. Russ. J. Herpetol., 17(1), 32. Fig. 26. Type locality of Agama pawlowskii Chernov et Dubinin, 1946 (red dot). Acanthosaura brachypoda Ananjeva, Orlov, Nguyen et Ryabov, 2011. Russ. J. Herpetol., 18(3), 196. and Shri Lanka: Cophotis dumbara V. A. M. P. K. Samarawickrama, Ranawana, Rajapaksha, Ananjeva, Orlov, Ranasinghe, Samarawaickrama et V. A. P. Samarawickrama, 2006. Russ. J. Herpetol., 10(1), 34. In the 1990s a joint complex study of phylogeny and biogeography of agamid lizards was developed in cooperation with Theodore J. Papenfuss and J. Robert Macey (Museum of Vertebrate Zoology, University of California, Berkeley, Department of Biology, Washington University, St. Louis, USA, and Department of Evolutionary Genomics, Joint Genome Institute, California, USA) (Macey et al., 1998, 2000a, b, Macey, Ananjeva, 2004). U. Joger and N. B. Ananjeva worked together on the project Molecular Systematics and phylogeny of the arboreal agamid lizards genus Acanthosaura (Grey, 1831) supported by a grant received from the corporation Volkswagen. This project, as well as a study of systematics and variability of mountain rock agamas of «Laudakia caucasia» complex was fulfilled with participation of the late young researcher Svetlana Kalyabina-Hauf (Kalyabina et al., 2004; Ananjeva and Kalyabina-Hauf, 2006). In 2003 Ananjeva and Milto collected data and samples together with Australian herpetologists Jane Melville (Victoria Museum,, Melbourne, Australia) and Nick Clemann (Arthur Rylah Inst. for Environmental Research, Dept. of Sustainability and Environment, Heidelberg, Australia) in Central Asia which led to interesting results in ecology and phylogeny of agamid genera Trapelus and Phrynocephalus (Clemann et al., 2008; Melville et al., 2009). For many years, several projects on the morphology of integument and its derivatives (receptors, glandular

Research on Asian Agamid Lizards: Contribution of Zoological Institute, Russian Academy of Sciences 97 structures) of iguanomorph lizards and, in particular, agamids were developed in cooperation with a research team from the Laboratory of Ornithology and Herpetology, Institute of Zoology, Ministry of Education and Sciences, Almaty, Kazakhstan led by Tatyana Dujsebayeva (Ananjeva et al., 1991, 2001; Ananjeva and Dujsebayeva, 2006; Dujsebayeva et al., 2007, 2009; etc.). Participation in several projects on the morphology of the integument of agamids is reflected in the publication of papers and abstracts at the International Congress of Morphology in Bristol (1997). For several years, agamids have been the subject of joint studies with Ella M. Smirina (Institute of Developmental Biology, Moscow, Russia) of the aging and longevity of Palearctic agamids and in particular for verification of the hypothesis about the ephemerality of small species of desert lizards (Smirina and Ananjeva, 2001; Ananjeva et al., 2003). We also studied growth layers in different bones and acrodont teeth of agamid lizards (Smirina and Ananjeva, 2007) using the fact that agamid lizards are an exception among reptiles in contrast to other reptiles, in agamids tooth replacement is restricted to the anterior pleurodont teeth only, but postcanine acrodont teeth are not replaced during their lifetime. Agamids of the genus Laudakia were the subject of a joint study with herpetologists of Armenia, Mongolia, and Turkmenistan; recently a comprehensive morphological revision was prepared in cooperation with scientists from Zoologisches Forschungsmuseum A. Koenig, Bonn, Germany Wolfgang Böhme and Philipp Wagner and late Pakistan herpetologists Khalid Javed Baig (Baig et al., 2012). At the current stage of modern history in the Department of Herpetology Natalia Ananjeva, Daniel Melnikov, and Konstantin Milto and associate researcher Andrej Barabanov continue to study systematics and phylogeny of agamids (Barabanov and Ananjeva, 2007; Melnikov et al., 2008; Ananjeva, 2004, 2011; Ananjeva et al., 2006, 2008a, 2008b; Milto and Barabanov, 2012). Daniel Melnikov focused on morphological and molecular study of agamids of genera Phrynocephalus, Trapelus, and Pseudotrapelus (Melnikov and Ananjeva, 2010; Melnikov et al., 2012) and mites of the family Pterygosomatidae (Acari: Prostigmata) permanent parasites of lizards, and perspectives of their implication in inference of host phylogeny (Melnikov and Bochkov, 2010). Joint study of agamids is conducted through joint projects with Yuezhao Wang and Xianguang Guo (Chengdu Institute of Biology, Chinese Academy of Sciences, China) (Ananjeva, Wang, 2008; Ananjeva et al., 2011), Nguyen Quang Truong, Institute of Ecology and Biological Resources, and Tao Thien Nguyen, Vietnam National Museum of Nature, Vietnamese Academy of Sciences and Technology, Salvador Carranza (Institut de Biologia Evolutiva (CSIC-UPF), Barcelona, Spain) and Jingzhong Fu (University of Guelph, Canada). Acknowledgments. The authors are sincerely grateful to the researchers of the Zoological Institute, RAS Helen V. Dubinina, Andrej V. Barabanov, Igor V. Doronin, Daniel A. Melnikov, Konstantin D. Milto, and Nikolai L. Orlov for their helpful assistance in preparation of this manuscript. Steven C. Anderson kindly provided linguistic editorial comments and corrections of English. 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