蜥蜴血液寄生蟲的多樣性是一個尚待探查的領域。過去的研究多僅靠蜥蜴血液抹片呈現的顯微影像鑑定物種，然而血液抹片所能提供的寄生蟲形態特徵相當有限，且僅包含一個生活史階段的特徵，易造成誤判與同物異名（synonyms）的現象。另外，顯微影像技術在低感染強度（intensity）的個體中，較易產生偽陰性（false-negative）結果。近年分子技術的開發，提供了另一種有效檢測血液寄生蟲的方法。本研究發現在台東鯉魚山公園、蘭嶼及綠島等地的斯文豪氏攀木蜥蜴身上，至少存在兩種分屬頂複器門（Apicomplexa）和線蟲門（Nematoda）的血液寄生蟲，由於過去在台灣並沒有相關的研究資料，因此，本研究除了蒐集血液寄生蟲的基礎生態與分子資料外，欲檢測以下問題：（1）所發現的血液寄生蟲其近源類群為何？（2）分子技術與顯微影像技術檢測血液寄生蟲的何者有較高的敏感度？（3）在不同地區所發現的血液寄生蟲是否分屬不同物種？（4）何時是血液寄生蟲盛行率最高的季節？（5）寄生蟲感染與哪些宿主的特徵有關？本研究收蒐集了不同季節6個地點共1,196隻攀蜥，藉由血液抹片、分子鑑定技術與親緣關係重建技術（phylogenetic reconstruction），確認所感染寄生蟲分別為巴氏簇蟲屬（Bartazoon spp.）與盤尾絲蟲科（Onchocercidae spp.）物種。而在形態資料比較上顯示兩種血液寄生
蟲在不同島上皆有顯著差異，然而，仍有待進一步的生活史資料證實，是否分屬為不同物種。巴氏簇蟲（Bartazoon sp.）在春季有最高的盛行率（6%）而微血絲蟲（microfilaria）則在秋季有最高的盛行率（16%）。相對感染風險（Relative Risk）的結果顯示，在蘭嶼的成蜥與幼蜥相比有顯著較高的感染風險（約21倍；95% CI: 7-68）。本研究提供台灣蜥蜴血液寄生蟲首次的發現記載，並成功開發巴氏簇蟲（Bartazoon sp.）的高檢測敏感度（sensitivity）的專一性引子。

Research of saurian blood parasite diversity represents a rather unexplored field. Definition of new species in previous studies based mainly on direct observation on reptile blood smears. However, specimens with low infection intensity could result to false-negative during the evaluation process. In recent years, molecular technique provides novel detection methods with higher precision. In this research, I collected 1,196 lizard specimens from 6 population in different seasons, and recorded at least two different phyla (Apicomplexa and Nematoda) of hemoparasites in Japalura swinhonis from Taitung Liyushan Park, Green Island and Orchid Island. Observation on blood smears, PCR assays and phylogenetic reconstruction indicated that the two blood parasites are close related to Bartazoon spp. and Onchocercidae spp. respectively. Both parasites, Bartazoon spp. and microfilaria, represent genetic similarity but morphological difference between two islands. Bartazoon spp. represent the highest prevalence in Spring (6%), while microfilaria in Autumn (16%). Evaluation on Relative Risk showed that adult lizards had 21 times (95% CI: 7-68) risk of infection by blood parasites compared to juveniles in Orchid Island. This research provides the first records of lizard blood parasites in Taiwan, and successfully improve the detection efficiency of Bartazoon spp. by using molecular techniques.

Alicata JE. A new genus for the nematode Filaria cistudinis Leidy, 1856, of the family Filariidae. J Wash Acad Sci. 1933;23(2):95-100.
Anderson RC. Nematode parasites of vertebrates: their development and transmission. 2nd ed. CABI Publishing: Wallingford, England; 2000. p. 467-532.
Bain O. Sulahian A. Trois nouvelles filaires du genre Oswaldofilaria chez des lézards sud-américains; essai de classification des Oswaldofilariinae. Bulletin du Muséum National d’Histoire Naturelle. 1974;156:827-841.
Bain O, Baldwin JG, Beveridge I, Bezerra TC, Braeckman U, Coomans A, Decraemer W, Derycke S, Durette-Desset M-C, Fonseca G. Handbook of Zoology. Gastrotricha, Cycloneuralia and Gnathifera Volume 2: Nematoda, In: Schmidt-Rhaesa A, editors. Walter de Gruyter; 2013.
Banoo S, Bell D, Bossuyt P, Herring A, Mabey D, Poole F, Smith PG, Sriram N, Wongsrichanalai C, Linke R, O'Brien R, Perkins M, Cunningham J, Matsoso P, Nathanson CM, Olliaro P, Peeling RW, Ramsay A. Evaluation of diagnostic tests for infectious diseases: general principles. Nat Rev Microbiol. 2008;5:S16–S28.
Barta JR, Ogedengbe JD, Martin DS, Smith TG. Phylogenetic position of the adeleorinid coccidia (Myzozoa, Apicomplexa, Coccidia, Eucoccidiorida, Adeleorina) inferred using 18S rDNA sequences. J Eukaryot Microbiol. 2012;59(2):171-180.
Boulianne B, Evans RC, Smith TG. Phylogenetic analysis of Hepatozoon species (Apicomplexa: Adeleorina) infecting frogs of Nova Scotia, Canada, determined by ITS-1 sequences. J Parasitol. 2007;93(6):1435-1441.
Casiraghi M, Anderson TJC, Bandi C, Bazzocchi C, Genchi C. A phylogenetic analysis of filarial nematodes: comparison with the phylogeny of Wolbachia endosymbionts. Parasitology. 2001;122(01):93-103.
Casiraghi M, Bain O, Guerrero R, Martin C, Pocacqua V, Gardner SL, Franceschi A, Bandi C. Mapping the presence of Wolbachia pipientis on the phylogeny of filarial nematodes: evidence for symbiont loss during evolution. Int j parasitol. 2004;34(2):191-203.
Chabaud AG, Anderson RC. Brygoo ER. Cinq Filaires de Reptiles malgaches. Mémoires de l’Institut Scientifique de Madagascar. 1959;13(A):103-126.
Chen SF, Chen SL, Li ZL, Lin HC, Chang MH. A survey of terrestrial vertebrates on Green island. Journal of National Park. 2009;19(3):1-22.
Darriba D, Taboada GL, Doallo R, Posada D. jModelTest 2: more models, new heuristics and parallel computing. Nat Methods. 2012;9(8):772.
Davies AJ., and Johnston MRL. The biology of some intraerythrocytic parasites of fishes, amphibia and reptiles. Adv Parasitol. 2000;45:1-107.
Galvani AP. Age-dependent epidemiological patterns and strain diversity in helminth parasites. J Parasitol. 2005;91(1):24-30.
García‐ramírez A, Delgado‐garcía JD, Foronda‐rodríguez P, Abreu‐acosta N. Haematozoans, mites and body condition in the oceanic island lizard Gallotia atlantica (Peters and Doria, 1882) (Reptilia: Lacertidae). J Nat Hist. 2005;39(17):1299-1305.
Garrido M, Pérez-Mellado V. Prevalence and intensity of blood parasites in insular lizards. Zoologischer Anzeiger-A Journal of Comparative Zoology. 2013;252(4):588-592.
Goodwin MH, Tommy KS. The course of natural and induced infections of Plasmodium floridense Thompson and Huff in Sceloporus undulatus undulatus (Latreille). Am J Trop Med Hyg. 1952;1(5):773-783.
Guindon S, Dufayard JF, Lefort V, Anisimova M, Hordijk W, Gascuel O. New algorithms and methods to estimate maximum-likelihood phylogenies: assessing the performance of PhyML 3.0. Syst biol. 2010;59(3):307-321.
Guindon S, Gascuel O. A simple, fast and accurate method to estimate large phylogenies by maximum-likelihood. Syst biol. 2003;52:696-704.
Gutiérrez-López R, Gangoso L, Martínez-de la Puente J, Fric J, López-López P, Mailleux M, Muñoz J, Touati L, Samraoui B, Figuerola, J. Low prevalence of blood parasites in a long-distance migratory raptor: the importance of host habitat. Parasit Vectors. 2015;8(1):1.
Hatcher MJ, Dick JT, Dunn AM. How parasites affect interactions between competitors and predators. Ecol Lett. 2006;9:1253–1271.
Hillis DM, Dixon MT. Ribosomal DNA: molecular evolution and phylogenetic inference. Q Rev Biol. 1991;411-453.
Honigberg BM. A contribution to systematics of the non-pigmented flagellates. In: Ludvík J, Lom J, Vávra J, editors. Progress in Protozoology: proceedings of the first International Congress on protozoology held at Prague. Academic Press. 1963.
Jeanmougin F, Thompson JD, Gouy M, Higgins DG, Gibson TJ. Multiple sequence alignment with Clustal X. Trends Biochem Sci. 1998;23:403-405.
Karadjian G, Chavatte JM, Landau I. Systematic revision of the adeleid haemogregarines, with creation of Bartazoon n. g., reassignment of Hepatozoon argantis Garnham, 1954 to Hemolivia, and molecular data on Hemolivia stellata. Ucla Sym Bi. 2015;22(31): 1-15.
Khanna RK. A new filarial worm from a North American snake. J Helminthol. 1933;11(2):105-108.
Knowles SCL, Wood MJ, Alves R, Wilkin T, Bensch S, Sheldon BC. Molecular epidemiology of malaria prevalence and parasitaemia in a wild bird population. Mol Ecol. 2011;20: 1062–1076.
Lefoulon E, Bain O, Bourret J, Junker K, Guerrero R, Cañizales I, Kuzmin Y, Satoto TBT, Cardenas-Callirgos JM, Lima SS, Raccurt C, Mutafchiev Y, Gavotte L, Martin C. Shaking the tree: multi-locus sequence typing usurps current onchocercid (Filarial Nematode) phylogeny. PLoS Negl Trop Dis. 2015;9(11):e0004233.
Leveille AN, Ogedengbe ME, Hafeez MA, Tu HH, Barta JR. The complete mitochondrial genome sequence of Hepatozoon catesbianae (Apicomplexa: Coccidia: Adeleorina), a blood parasite of the green frog, Lithobates (formerly Rana) clamitans. J Parasitol. 2014;100(5):651-656.
Levine ND. Taxonomy of the Sporozoa. J Parasitol. 1970;56(4):208-9
López-Neyra CR. Revision de la superfamilia Filarioidea (Weinland, 1858). Revtaibér. Parasit. 1956;1(6):3-225.
Macfie JWS. Saurositus agamae, ng, n. sp. a filarioid parasite of the lizard Agama colonorum. Ann Trop Med Parasitol. 1924;18(3):409-412.
Mader DR. Reptile medicine and surgery. 2nd ed. Elsevier Health Sciences; 2006. p. 360.
Maia, JP, Harris DJ, Carranza S, Gómez-Díaz E. A comparison of multiple methods for estimating parasitemia of hemogregarine hemoparasites (Apicomplexa: Adeleorina) and its application for studying infection in natural populations. PloS one. 2014a;9(4):e95010.
Maia JP, Crottini A, Harris DJ. Microscopic and molecular characterization of Hepatozoon domerguei (Apicomplexa) and Foleyella furcata (Nematoda) in wild endemic reptiles from Madagascar. Ucla Sym Bi. 2014b;21:47
Martinsen ES, Perkins SL, Schall JJ. A three-genome phylogeny of malaria parasites (Plasmodium and closely related genera): evolution of life-history traits and host switches. Mol Phylogenet Evol. 2008;47(1):261-273.
McLaren DJ. Ultrastructural studies on microfilariae (Nematoda: Filarioidea). Parasitology. 1972;65(2):317-332.
Meyer A, Todt C, Mikkelsen NT, Lieb B. Fast evolving 18S rRNA sequences from Solenogastres (Mollusca) resist standard PCR amplification and give new insights into mollusk substitution rate heterogeneity. BMC Evol. Biol. 2010;10(1):70.
Miyata A, Miyagi I, Tsukamoto M. Haemoprotozoa Detected from the Cold-blooded Animals in Ryukyu Islands. Tropical medicine. 1978;20(2):97-112.
Mullin SW. Gonofilaria rudnicki gen. et sp. n. (Nematoda: Filarioidea) from Malaysian lizards. Proc Helminthol Soc Wash. 1973;40: 282-285.
O’Dwyer LH, Moço TC, dos Santos Paduan K, Spenassatto C, da Silva RJ, Ribolla PEM. Description of three new species of Hepatozoon (Apicomplexa, Hepatozoidae) from Rattlesnakes (Crotalus durissus terrificus) based on molecular, morphometric and morphologic characters. Exp Parasitol. 2013;135(2):200-207.
Pandit CG, Pandit SR, SEETHARAMA IYER PV. The Development of the Filaría Conispiculum guindiensis (1929) in C. fatigans, with a Note on the Transmission of the Infection. Indian J Med Res. 1929;17(2):421-429.
Perkins SL, Keller AK. Phylogeny of nuclear small subunit rRNA genes of hemogregarines amplified with specific primers. J Parasitol. 2001;87(4):870-876.
Piersma T. Do global patterns of habitat use and migration strategies co-evolve with relative investments in immunocompetence due to spatial variation in parasite pressure? Oikos. 1997;80:623–31.
Poulin R. Variation in infection parameters among populations within parasite species: intrinsic properties versus local factors. Int J Parasitol. 2006a;36:877–885.
Poulin R. Evolutionary ecology of parasites. Princeton university press. 2006b.
Potts FA, Borradaile LA. The phylum Nematoda. In: Eastham LES, Saunders JT, editors. The Invertebrata, a manual for the use of students. Cambridge Univ. Press; 1932. p. 214-227.
Rambaut A, Suchard M, Drummond A. 2013. Tracer v1. 6: An MCMC Trace Analysis Tool. Institute of Evolutionary Biology, University of Edinburgh. <http://tree.bio.ed.ac.uk/software/tracer/>.
Ronquist F, Teslenko M, van der Mark P, Ayres DL, Darling A, Höhna S, Larget B, Liu L, Suchard MA, Huelsenbeck, JP. MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic biology. 2012;61(3):539-542.
Sabagh LT, Borges-Júnior V, Winck G, Viana L, Rocha C. Low prevalence of hemoparasites in a lizard assemblage from a coastal environment in southeastern Brazil. Herpetology Notes. 2015;8:413-416.
Sato H, Takano A, Kawabata H, Une Y, Watanabe H, Mukhtar MM. Trypanosoma cf. varani in an imported ball python (Python reginus) from Ghana. J Parasitol. 2009;95(4):1029-1033.
Schall JJ, Albert FB, Robert WP. Lizards infected with malaria: physiological and behavioral consequences. Science. 1982;217(4564):1057-1059.
Schall JJ. Parasite-mediated competition in Anolis lizards. Oecologia. 1992;92(1):58-64.
Schall JJ. Malarial parasites of lizards: Diversity and ecology. Adv Parasitol. 1996;37:255-333.
Scorza JV. Anaemia in lizard malaria infections, Parassitologia. 1971;13:391-405.
Seurat LG. Filaires des Reptiles et des Batraciens. Bulletin de la Société d’Histoire Naturelle d’Afrique du Nord. 1917;8:236–242.
Sloboda M, Kamler M, Bulantová J, Votýpka J, Modrý D. A new species of Hepatozoon (Apicomplexa: Adeleorina) from Python regius (Serpentes: Pythonidae) and its experimental transmission by a mosquito vector. J Parasitol. 2007;93(5):1189-1198.
Smith AJ. A new filarial species (F. mitchelli n. sp.) found in Heloderma suspectum and its larvae in a tick parasitic upon the gila monster. Univ Pennsylvania Med Bull. 1910;23:487-497.
Smith TG. The genus Hepatozoon (Apicomplexa: Adeleina). J Parasitol. 1996;82(4):565-585.
Sulahian A, Schacher JF. Brygoofilaria agamae gen. et sp. n. (Nematoda: Filarioidea) from the lizard Agama stellio in Lebanon. J Parasitol. 1968;831-833.
Super PE, van Riper IIIC. A comparison of avian hematozoan epizootiology in two California coastal scrub communities. J Wildl Dis. 1995;31:447–61.
Svahn K. Incidence of blood parasites of the genus Karyolysus (Coccidia) in Scandinavian lizards. Oikos. 1974:43-53.
Tamura K, Stecher G, Peterson D, Filipski A, Kumar S. MEGA6: molecular evolutionary genetics analysis version 6.0. Mol. biol. evol. 2013;30(12):2725-2729.
Telford SR. Interpopulation variation of a saurian malaria, Plasmodium sasai telford & ball, 1969, in three host species distributed within a range of 24° north latitude. Int J Parasitol. 1982;12(1):17–22.
Telford SR. Hemoparasites of the Reptilia: Color Atlas and Text. CRC Press; 2009.
Travassos L. Sobre os filarideos dos crocodillos sul-americanos. Memórias do Instituto Oswaldo Cruz. 1933;27(2):159-164.
Uetz P, Jirí Hošek (eds.). The Reptile Database, http://www.reptile-database.org accessed 2016.
Ujvari B, Madsen T, Olsson M. High prevalence of Hepatozoon spp.(Apicomplexa, Hepatozoidae) infection in water pythons (Liasis fuscus) from tropical Australia. J Parasitol. 2004;90(3):670-672.
van As J. Ecology, taxonomy and possible life cycles of blood protozoans infecting crag lizards (Pseudocordylus spp.) from the eastern Free State highlands. 2012. PhD Thesis. University of the Free State.