中文摘要
草蜥屬（Takydromus）在分類上屬於有鱗目（Squamata）、蜥蜴科（Lacertidae），主要分佈於東亞地區，喜歡棲息於草生地或森林邊緣。這個屬在本研究進行之初共有16至17種，並依其型態特徵區分為草蜥（Takydromus）與地蜥（Platyplacopus）兩個亞屬。這個屬有大約一半的種類分佈於日本、琉球與台灣，並形成當地的島嶼特有種。因此，利用分子工具，並結合地質變動的歷史，從不同的分類階層探討草蜥屬在東亞島嶼地區的演化歷史，即為本論文的主要目的（第一章）。
首先藉由定序完整的粒線體12S rRNA基因，以建立草蜥屬的親緣關係（第二章）。分子親緣關係顯示草蜥屬可粗略分為三大群，但是草蜥亞屬與地蜥亞屬並不各自形成單系群，而且部分種類的演化地位與傳統外部型態推測的結果有明顯的差異。草蜥屬在東亞島嶼地區的演化過程與島嶼之間相連接的歷史具有密切的關連，而島嶼之間因為冰河退去所造成的隔離演化效應（vicariance），可能是造成島嶼特有草蜥發生種化的主要因素。另一方面，由於草蜥種與種之間的遺傳距離上下差距極大，顯示較原始草蜥與較晚近草蜥的種化事件並非源自於同一系列的地質歷史事件。因此，依據分子生物與地質變動歷史的資料，我提出草蜥屬在東亞地區的「多次入侵」理論，用以解釋這些不同群草蜥在台灣的分佈及起源。根據分子時鐘的估算，可以讓我們大致掌握歷史上每一個分歧事件發生的時間。這個估算出來的演化速率，亦將在後續的討論之中多次被引用。
島嶼間的隔離效應雖然可以解釋大部分草蜥的種化機制，但是卻不含括所有可能的原因。台灣島由於地形崎嶇，山脈本身對陸域生物形成明顯的隔離效應。在第三章之中，我要介紹草蜥島內種化（in situ speciation）的案例。我於全島各地廣泛採集台灣草蜥（T. formosanus）的樣本，定序了兩個粒線體基因片段之後，發現東部、西部與北部的台灣草蜥在序列上擁有極大的差異，在外觀上亦具備可鑑別的特徵。其中，北部的族群與東部的族群應可提升為獨立的新種。這三個種與高海拔的雪山草蜥（T. hsuehshanensis）、高緯度的白條草蜥（T. wolteri）共同形成一個近緣的「台灣草蜥複合種群」（T. formosanus species complex），而這五個種之間的演化過程，亦將在第三章中進行詳細的討論。
在第四章，探討對象則鎖定為單一種內的演化歷史。我採集來自台灣西部14個地點的蓬萊草蜥（T. stejnegeri），進行族群遺傳與親緣地理學的探討。依據分子證據，蓬萊草蜥在種下可分為二到三個明確的系群（lineage），而其中最主要的兩個系群在台灣北部形成共域分佈的交會帶。我因此推測蓬萊草蜥極有可能在冰河時期居於不同的避難所內，因而形成明顯系群的分化；並由於冰河時期結束之後產生明顯的族群擴張，因此造成兩個系群在分佈上的重疊。利用網狀聚群分析（nested clade analysis）、中性檢測（neutrality tests）與單一位點突變頻率分析（frequency spectrum）等不同方法，結果均顯示蓬萊草蜥在最近地質變動歷史中的族群擴張現象。
由於冰河事件在地質史上的反覆出現，導致所有的陸域生物均受到這種氣候反覆變化的影響，進而發生族群數量的擴張與退縮。我將網狀聚群分析（nested clade analysis）的技術應用在台灣草蜥複合種群的四個特有種上，試圖找出物種在族群波動歷史上的共通性（第五章）。結果顯示分佈於平地的北部新種與東部新種的族群遺傳結構與蓬萊草蜥類似，並顯現出更明顯的族群歷史性波動。這三個種均呈現明確但部分重疊的兩個系群，極為可能是冰河時期結束之後發生族群擴張的結果。但是分佈於西部平地的台灣草蜥族群，相較之下呈現較為穩定的族群歷史。我推測其與共域之蓬萊草蜥間的種間競爭，可能是導致其族群在冰河結束之後無法大量擴張的因素。另一方面，依據推測，分佈於高山的雪山草蜥，應該在冰河結束之後呈現族群退縮的趨勢。但可能受限於網狀聚群分析的分析效力，亦可能受限於樣本數量的大小，這個結果在分析之中並不明顯。
在經過這些研究之後，發現台灣地區同時分佈著七種草蜥，並包含六個特有種。由於草蜥屬的共域分佈現象在其他地方相當罕見，因此這樣的種類密度，在世界上是獨一無二的。然而直至目前為止，我們對草蜥生態上或分佈上的瞭解，仍然極為有限。在第六章之中，我將對未來的相關研究題材提出展望，並對上述的研究進行總結。

Abstract
The East Asian grass lizard genus, Takydromus Daudin in the family Lacertidae, is distributed in the Oriental and Palearctic regions. With specialized slender body shape and extraordinarily long tails, most Takydromus species inhabit in grasslands, dense scrub, or forest edge. At the beginning of my study in 1998, there were 16 or 17 species recognized in this genus, belonging to two subgenera: Takydromus and Platyplacopus. One of the most spectacular features of grass lizards in biogeography is the high percentage of endemism in the islands. Almost half of Takydromus species are endemic to the islands aligned along the Pacific coast of the East Asian continent. The formation and evolution of Takydromus in this region is an interesting issue in biogeography research (Chapter 1).
In Chapter 2, I sequenced the complete mitochondrial 12S rRNA from 14 Takydromus OTUs. Phylogenetic relationship was constructed by MP, NJ and ML methods, and the results of these three criterions are congruent. Three major groups were identified within this genus, whereas the monophyly of both subgenera was not supported. According to our deduction, the speciation process of Takydromus is closely related to the pattern of land connection among island groups influenced by glaciations. The vicariance between neighboring islands triggered the divergence between species. Furthermore, a model of multiple colonizations was proposed to explain the prominent difference of interspecific distance among different Takydromus groups. A rough estimation on the evolutionary rate helps us to deduce the age of each historical event, while this estimation would also be applied in the following sections.
Although the model of inter-island vicariance explained the distribution of most Takydromus species, an “ in situ speciation ” model was further proposed to explain the biodiversity of grass lizard in Taiwan. In Chapter 3, I collected T. formosanus from 27 locations throughout the entire island. Mitochondrial sequencing represented significant differentiation among eastern, western, and northern populations in Taiwan. With diagnostic morphological characters, two of these “evolutionarily significant units”, the northern lineage and the eastern lineage, were suggested to evaluate as new species. The combination of T. formosanus, T. hsuehshanensis, and T. wolteri formed a monophyletic group, and was defined as “Takydromus formosanus species complex”. The evolutionary relationship among the five members within this species complex was discussed in Chapter 3.
In Chapter 4, I focused on the population genetic study of Takydromus stejnegeri, another island-endemic species in Taiwan. Fourteen populations were collected, sequenced, and analyzed with nested clade analysis (NCA). Two to three major lineages were detected in T. stejnegeri, with partially overlapping distribution in northern Taiwan. Their current distribution was deduced to be a post-glacial expansion from two or three refugial regions. The recent history of “range expansion” for T. stejnegeri was supported by the evidence from NCA, neutrality tests, and frequency spectrum.
The periodical occurrence of glaciations caused the demographic fluctuation of these lizards. To figure out the pattern of such influences, I applied the analysis of NCA to the four endemic units of T. formosanus species complex in Chapter 5. The genetic pattern of the northern species (Takydromus sp. N) and the eastern species (Takydromus sp. E) were similar to that in T. stejnegeri, revealing a deep intraspecific branching with partially overlapping distribution. The demographic fluctuation in these two species was prominent, but not present in T. formosanus distributed in western Taiwan. An interspecific competition with T. stejnegeri is a probable explanation for its comparatively stable demographic history. On the other hand, the expected population decline in T. hsuehshanensis was not observed, probably owing to the analytical power of NCA or the adequate sampling for this species.
With seven species including six endemics, the island of Taiwan represents the highest biodiversity of Takydromus in the world. However, our knowledge on these secretive lizards is still limited. In Chapter 6, I will make some prospect on concerning studies in the future, and make a conclusive comment on the study of Takydromus in the past five years.

References
Arnold, E. N. 1989. Towards a phylogeny and biogeography of the Lacertidae: relationships within an Old-World family of lizards derived from morphology. Bull. Br. Mus. Nat. Hist. (Zool.) 55: 209-257.
Arnold, E. N. 1997. Interrelationships and evolution of the east Asian grass lizards, Takydromus (Squamata: Lacertidae). Zool. Jour. Linn. Soc. 119: 267-296.
Avise, J. C. 2000. Phylogeography - the history and formation of species. Harvard University Press. London, England.
Avise, J. C., and D. Walker. 1998. Pleistocene phylogeographic effects on avian populations and the speciation process. Proc. Roy. Soc. London 265: 457-463.
Bogadek, A., and M. W. N. Lau. 1997. A revised checklist of Hong Kong amphibians and reptiles. Mem. Hong Kong Nat. Hist. Soc. 21: 173-188.
Boulenger, G. A. 1894. Descriptions of a new lizard and a new fish obtained in Formosa by Mr. Holst. Ann. Mag. Nat. Hist. 6: 462-463.
Boulenger, G. A. 1917. A revision of the lizards of the genus Takydromus. Mem. Asiatic Soc. Bangal 5: 207-235.
Brown, W. M., M. George, and A. C. Wilson. 1979. Rapid evolution of animal mitochondrial DNA. Proc. Natl. Acad. Sci. USA 76: 1967-1971.
Chen, I.-S., and L.-S. Fang. 1999. The freshwater and estuarine fishes of Taiwan. National Museum of Marine Biology and Aquarium, Taiwan.
Chi, W. R., J. Namson, and J. Suppe. 1981. Stratigraphic record of plate interactions in Coastal Range, eastern Taiwan. Mem. Geol. Soc. China 4: 155-194.
Chou, W.-H., N. Q. Truong, and O. Pauwels. 2001. A new species of Takydromus (Reptilia: Lacertidae) from Vietnam. Herpetologica 57: 497-508.
Clement, M., D. Posada, and K. A. Crandall. 2000. TCS: a computer program to estimate gene genealogies. Mol. Ecol. 9: 1657-1659.
Daudin, F. M. 1802. Histoire Naturelle des Reptiles, vol. 3. F. Dufart, Paris.
De Peer, Y. V., I. V. Den Broeck, P. de Rijk, and R. de Wachter. 1994. Database on the structure of small ribosomal subunit RNA. Nucleic Acids Res. 22: 3488-3494.
Fang, Y.-P. 2001. Systematics and phylogeographic studies of the genus Crocidura in Taiwan (Insectivora: Soricidae). Ph. D. Thesis.of National Taiwan University. Taipei, Taiwan.
Fu, J. 1998. Toward the phylogeny of the family Lacertidae: implications from mitochondrial DNA 12S and 16S gene sequences (Reptilia: Squamata). Mol. Phylog. Evol. 9: 118-130.
Fu, J. 2000. Toward the phylogeny of the family Lacertidae - Why 4708 base pairs of mtDNA sequences cannot draw the picture. Biol. Jour. Linn. Soc. 71: 203-217.
Fu, Y.-X., and W.-H. Li. 1993. Statistical tests of neutrality of mutations. Genetics 133: 693-709.
Fuller, S., P. Baverstock, and D. King. 1998. Biogeographic origins of goannas (Varanidae): a molecular perspective. Mol. Phylog. Evol. 9: 294-307.
Hachisuka, M., and T. Udagawa. 1950. Contributions to the ornithology of Formosa, Part 1. Quart. Jour. Taiwan Mus. 3: 187-280.
Harris, D. J., E. N. Arnold, and R. H. Thomas. 1998. Relationships of lacertid lizards (Reptilia: Lacertidae) estimated from mitochondrial DNA sequences and morphology. Proc. R. Soc. Lond. 265: 1939-1948.
Harrison, R. G. 1998. Linking evolutionary pattern and process. In Endless forms - species and speciation. Oxford University Press, Inc. New York.
Hickson, R. E., C. Simon, A. Cooper, G. S. Spicer, J. Sullivan, and D. Penny. 1996. Conserved sequence motifs, alignment, and secondary structure for the third domain of animal 12S rRNA. Mol. Biol. Evol. 13: 150-169.
Hikida, T. 1993. Phylogenetic relationships of the skinks of the genus Eumeces (Scincidae: Reptilia) from East Asia. Jpn. Jour. Herpetol. 15: 1-21.
Hikida, T., and J. Motokawa. 1999. Phylogeographical relationships of the skinks of the genus Eumeces (Reptilia: Scincidae) in East Asia. In Tropical Island Herpetofauna - Origin, Current Diversity, and Conservation (H. Ota, Ed.), pp. 231-247. Elsevier Science B.V., Amsterdam.
Hikida, T., and H. Ota. 1997. Biogeography of reptiles in the subtropical East Asian Islands. The Symposium on the Phylogeny, Biogeography and Conservation of Fauna and Flora of East Asian Region, Taipei, Taiwan, ROC, pp. 11-18.
Ho, C.-S. 1982. Tectonic evolution of Taiwan - Explanatory text of the tectonic map of Taiwan. The Ministry of Economic Affairs, Taiwan, R.O.C. Taipei, Taiwan.
Hsieh, B.-J. 1998. Phylogenetic study on the family Clausiliidae in Taiwan with morphological characters and mt-large (16S) rRNA. Master thesis of National Taiwan University. Taipei, Taiwan.
Hsu, Y.-T. 1999. The evolutionary relationship of Chinese bulbul (Pycnonotus sinensis) and Taiwan bulbul (P. taivanus) revealed by their population genetic structure. Master thesis of National Taiwan University. Taipei, Taiwan.
Huelsenbeck, J. P. and K. A. Crandall. 1997. Phylogeny estimation and hypothesis testing using maximum likelihood. Ann. Rev. Ecol. Syst. 28: 437-466.
Kimura, M. 1980. A simple method for estimating evolutionary rates rates of base substitutions through comparative studies of nucleotide sequences. Jour. Mol. Evol. 16: 111-120.
Kimura, M. 2000. Paleogeography of the Ryukyu Islands. Tropics 10: 5-24.
Kishino, H. and M. Hasegawa. 1989. Evaluation of the maximum likelihood estimate of the evolutionary tree topology from DNA sequence data, and the branching order of the Hominoidea. J. Mol. Evol. 29: 170-179.
Kizaki, K., and I. Oshiro. 1980. The origin of the Ryukyu Islands. In Natural History of the Ryukyus (K. Kizaki, Eds.), Tsukiji-shokan, Tokyo.
Klicka, J., and R. M. Zink. 1997. The importance of recent ice ages in speciation: a failed paradigm. Science 277: 1666-1669.
Kumar, S., K. Tamura, I. B. Jakobsen, and M. Nei. 2001. MEGA2: Molecular Evolutionary Genetics Analysis. Arizona State University, Tempe, Arizona, USA.
Liew, P.-M. 1994. Palynological study of Quaternary lake deposits in Yuan-Shan, Ilan. NSC82-0202-M002-113. National Science Council, Taipei, Taiwan.
Lin, J.-Y., and H.-Y. Cheng. 1981. A new species of Takydromus (Sauria: Lacertidae) from Taiwan. Bull. Inst. Zool. Acad. Sinica 20: 43-47.
Lin, S.-M., C. A. Chen, and K.-Y. Lue. 2001. Molecular Phylogeny and Biogeography of the Grass Lizards Genus Takydromus (Reptilia: Lacertidae) of Eastern Asia. The 2001 Joint Annual Meetings of the Society for the Study of Amphibians and Reptiles and the Herpetologists' League. Indianapolis, Indiana, USA.
Lin, S.-M., C. A. Chen, and K.-Y. Lue. 2002. Molecular Phylogeny and Biogeography of the Grass Lizards Genus Takydromus (Reptilia: Lacertidae) of East Asia. Mol. Phylogenet. Evol. 22: 276-288.
Lin, S.-M., and K.-Y. Lue. 2000. Phylogeography of the Takydromus lizard on East Asian islands: evidence based on mitochondrial 12S rRNA sequence. Fourth Asian Herpetological Conference. Chengdu, China.
Lue, K.-Y., and J.-S. Lai. 1991. Wildlife Data Bank of Taiwan (3) Lizards (I). Council of Agriculture, Taiwan. Taipei, Taiwan.
Lue, K.-Y., M.-C. Tu, and G. Shang. 1999. A Field Guide to the Amphibians and Reptiles of Taiwan.SWAN Pub. Co. Taipei, Taiwan.
MacArthur, R. H., and E. O. Wilson. 1966. The theory of island biogeography. Princeton University Press. Princeton, N.J., U.S.A.
Muse, S.V. and B. S. Weir. 1992. Testing for equality of evolutionary rates. Genetics 132: 269-276.
Neefs, J.-M., Y. V. De Peer, P. De Rijk, S. Chapelle, and R. De Wachter. 1993. Compilation of small ribosomal subunit RNA structure. Nucleic Acids Res. 21: 3025-3049.
Nixon, K. C., and Q. D. Wheeler. 1990. An amplification of the phylogenetic species concept. Cladistics 6: 211-223.
Ota, H. 1997. Historical biogeographical implications in the variation and diversity of amphibians and reptiles in Taiwan. The Symposium on the Phylogeny, Biogeography and Conservation of Fauna and Flora of East Asian Region. Taipei, Taiwan, ROC.
Ota, H. 1998. Geographic patterns of endemism and speciation in amphibians and reptiles of the Ryukyu Archipelago, Japan, with special reference to their paleogeographical implications. Res. Popul. Ecol. 40: 189-204.
Ota, H. 2000. The current geographic faunal pattern of reptiles and amphibians of the Ryukyu Archipelago and adjacent regions. Tropics 10: 51-62.
Ota, H., M. Honda, S.-L. Chen, T. Hikida, S. Panha, H.-S. Oh, and M. Matsui. 2002. Phylogenetic relationships, taxonomy, character evolution and biogeography of the lacertid lizards of the genus Takydromus (Reptilia: Squamata): a molecular perspective. Biol. Jour. Linn. Soc. 76: 493-509.
Ota, H., M. W. Lau, and A. Bogadek. 1994. Taxonomic status of the Hong Kong populations of the Hemidactylus garnotii-vietnamensis complex (Gekkonidae: Reptilia). Zool. Stud. 34: 126-130.
Otsuka, H., and A. Takahashi. 2000. Pleistocene vertebrate faunas in the Ryukyu Islands: their migration and extinction. Tropics 10: 25-40.
Pope, C. H. 1935. The Reptiles of China. Turtles, Crocodilians, Snakes, Lizards. American Museum of Natural History, New York.
Posada, D., and K. A. Crandall. 1998. Modeltest: testing the model of DNA substitution. Bioinformatics 14: 817-818.
Posada D., K. A. Crandall, and A. R. Templeton. 2000. GeoDis: A program for the cladistic nested analysis of the geographical distribution of genetic haplotypes. Mol. Ecol. 9:487-488.
Queiroz, K. d. 1998. The general lineage concept of species, species criteria, and the process of speciation. In Endless forms - species and speciation. Oxford University Press, Inc. New York.
Richards, C. M., and W. S. Moore. 1996. A phylogeny for the African treefrog family Hyperoliidae based on mitochondrial rDNA. Mol. Phylogenet. Evol. 5: 522-532.
Rozas, J., and R. Rozas. 1999. DNA Sequence Polymorphism Version 3.14.3. Universitat de Barcelona. Barcelona, Spain.
Schmidt, H. A., K. Strimmer, M. Vingron, and A. von Haeseler. 2002. TREE-PUZZLE 5.1: Maximum likelihood analysis for nucleotide, amino acid, and two-state data. Available from http://www.tree-puzzle.de/
Shen, S.-C. 1993. Fishes of Taiwan. Department of Zoology, National Taiwan University. Taipei, Taiwan.
Simonsen, K. L., G. A. Churchill, and C. F. Aquadro. 1995. Properties of statistical tests of neutrality for DNA polymorphism data. Genetics 141: 413-429.
Springer, M. S., L. J. Hollar, and A. Burk. 1995. Compensatory substitutions and the evolution of the mitochondrial 12S rRNA gene in mammals. Mol. Biol. Evol. 12: 1138-1150.
Suzuki, H., K. Tsuchiya, and N. Takezak. 2000. A molecular phylogenetic framework for the Ryukyu endemic rodents Tokudaia osimensis and Diplothrix legata. Mol. Phylogenet. Evol. 15: 15-24.
Swofford, D.L. 2001. PAUP 4.06b: Phylogenetic Analysis Using Parsimony and Other Methods. Sinauer Associates. Sunderland, Massachusetts.
Swofford, D. L. 2002. PAUP 4.0b10: Phylogenetic Analysis Using Parsimony and Other Methods. Sinauer Associates. Sunderland, Massachusetts.
Tajima, F. 1989a. The effect of change in population size on DNA polymorphism. Genetics 123: 597-601.
Tajima, F. 1989b. Statistical method for testing the neutral mutation hypothesis by DNA polymorphism. Genetics 123: 585-595.
Takeda, N., and H. Ota. 1996. Description of a new species of Takydromus from the Ryukyu Archipelago, Japan, and a taxonomic redefinition of T. smaragdinus Boulenger 1887 (Reptilia: Lacertidae). Herpetologica 52: 77-88.
Tamura, K., and M. Nei. 1993. Estimation of the number of nucleotide substitution in the control region of mitochondrial DNA in humans and chimpanzees. Mol. Biol. Evol. 10: 512-526
Templeton, A. R. 1983. Phylogenetic inference from restriction endonucleotide cleavage site maps with particular reference to the evolution of humans and apes. Evolution 37: 221-244.
Templeton, A. R. 1998. Nested clade analyses of phylogeographic data: testing hypotheses about gene flow and population history. Mol. Ecol. 7: 381-397.
Templeton, A. R., E. Boerwinkle, and C. F. Sing. 1987. A cladistic analysis of phenotypic associations with haplotypes inferred from restriction endonuclease mapping. I. Basic theory and an analysis of alcohol dehydrogenase activity in Drosophila. Genetics 117: 343-351.
Templeton, A. R., E. Routman, and C. A. Phillips. 1995. Separating population structure from population history: a cladistic analysis of the geographical distribution of mitochondrial DNA haplotypes in the tiger salamander, Ambystoma tigrinum. Genetics 140: 767-782.
Templeton, A. R., and C. F. Sing. 1993. A cladistic analysis of phenotypic associations with haplotypes inferred from restriction endonuclease mapping. IV. Nested analyses with cladogram uncertainty and recombination. Genetics 134: 659-669.
Toda, M. 1999. Historical biogeography of East Asian populations of Rana limnocharis (Amphibia: Anura): A review. In Tropical Island Herpetofauna - Origin, Current Diversity, and Conservation. Elsevier Science B.V. Amsterdam, the Netherlands.
Toda, M., M. Nishida, M. Matsui, K. Y. Lue, and H. Ota. 1998. Genetic variation among populations of the Indian rice frog, Rana limnocharis (Amphibia: Anura) within Taiwan: revealed by allozyme data. Herpetologica 54: 73-82.
Toda, M., M. Nishida, M. Matsui, G.-F. Wu, and H. Ota. 1997. Allozyme variation among East Asian populations of the Indian rice frog, Rana limnocharis (Amphibia: Anura). Bioch. Syst. Ecol. 25: 143-159.
Toda, M., M. Nishida, M.-C. Tu, T. Hikida, and H. Ota. 1999. Genetic variation, phylogeny and biogeography of the pit vipers of the genus Trimeresurus sensu lato (Reptilia: Viperidae) in the subtropical East Asian islands. In Tropical Island Herpetofauna - Origin, Current Diversity, and Conservation. Elsevier Science B.V. Amsterdam, the Netherlands.
Wang, H.-Y., M.-P. Tsai, M.-C. Tu, and S.-C. Lee. 2000. Universal primers for amplification of the complete mitochondrial 12S rRNA gene in vertebrates. Zool. Stud. 39: 61-66.
Webb, T. I., and P. J. Bartlein. 1992. Global changes during the last 3 million years: climate controls and biotic responses. Annu. Rev. Ecol. Syst. 23: 141-173.
Wise, C. A., M. Sraml, and S. Easteal. 1998. Departure from neutrality at the mitochondrial NADH dehydrogenase subunit 2 gene in humans, but not in chimpanzees. Genetics 148: 409-421.
Zhao, E.-M., and K. Adler. 1993. Herpetology of China.Society for the Study of Amphibains and Reptiles. Oxford, Ohio, USA.
Ziegler, T., and W. Bischoff. 1999. Takydromus (Platyplacopus) kuehnei vietnamensis ssp. n., eine neue Schnelläufereidechsen-Unterart aus Vietnam (Reptilia: Squamata: Lacertidae). Deutsche Gesellsch. Herpetol. Terrar. 35: 209-226.
Ziegler, T., W. Böhme, and W. Bischoff. 1999. Comments on the grass lizards (Lacertidae: Takydromus) of Vietnam and Myanmar. Hamadryad 24: 39-42.
Ziegler, T., H. X. Quang, and W. Böhme. 1998. Beitrag zur Kenntnis der Schnelläufer-Eidechsen Vietnams (Reptilia: Lacertidae: Takydromus). Herpetofauna 20: 24-34.