PROC. ENTOMOL. SOC. WASH. 107(2). 2005. pp. 245-253 THE TICKS (ACARI: IXODIDA: ARGASIDAE, IXODIDAE) OF TAIWAN: A SYNONYMIC CHECKLIST Richard G. Robbins Defense Pest Management Information Analysis Center, Armed Forces Pest Manage- ment Board, Walter Reed Army Medical Center, Washington, DC 20307-5001, U.S.A. (e-mail: richard.robbins@osd.mil) Abstract. — Eleven checklists of Taiwanese ticks, published or privately circulated be- tween 1935 and 1984, are compared with specimen records, chiefly in the U.S. National Tick Collection, yielding a list of 32 tick species thought to definitely occur in Taiwan and its adjacent islets: Argas pusillus Kohls, A. robertsi Hoogstraal, Kaiser & Kohls, Ornithodoros capensis Neumann, Amblyomma cordiferum Neumann, A. geoemydae (Can- tor), A. helvolum Koch, A. testudinarium Koch, Aponomma varanense (Supino), Boophihis microplus (Canestrini), Dermacentor taiwanensis Sugimoto, Haemaphysalis bandicota Hoogstraal & Kohls, H. canestrinii (Supino), H. doenitzi Warburton & Nuttall, H. flava Neumann, H. fonnosensis Neumann, H. hystricis Supino, H. kitaokai Hoogstraal, H. ma- ge shimaensis Saito & Hoogstraal, H. ornithophila Hoogstraal & Kohls, H. phasiana Saito, Hoogstraal & Wassef, H. taiwana Sugimoto, H. yeni Toumanoff, Ixodes acutitarsus (Karsch), /. granulatiis Supino, /. kuntzi Hoogstraal & Kohls, /. nipponensis Kitaoka & Saito, /. ovatus Neumann, /. persulcatus Schulze, /. simplex Neumann, /. vespertilionis Koch, Rhipicephalus haemaphysaloides (Supino), and R. sanguineus (Latreille). An ad- ditional 13 tick species previously believed to occur in Taiwan are excluded from this list. Key Words: ticks, Ixodida, checklist, synonymy, Taiwan At least 1 1 papers and reports intended consigned to the junior synonymy. With the to serve as checklists of the ticks of Taiwan recent emergence of Lyme bon-eliosis and (formerly Formosa) and vicinity were pub- human babesiosis in Taiwan (Shih and lished or privately circulated during the 20'^ Chao 1998, 1999; Shih et al. 1997. 1998). century (Kishida 1935, Schulze 1935, Ogu- it is imperative that these lists be reconciled ra 1936, Sugimoto 1939, Elishewitz 1943, with current taxonomic literature and a\ail- Anonymous 1944. Luh and Woo 1950. Shi- able specimen records to produce a single mada et al. 1961, Maa and Kuo 1966, Tseng reliable roster of the Taiwan tick fauna. 1978, and Hoogstraal letter no. 251. 14 Between 1955 and 1978. teams from February 1984, Smithsonian Institution Ar- U.S. Naval Medical Research Unit Number chives). However, when these lists are crit- Two (NAMRU-2), Taipei, scoured Taiwan ically compared, it soon becomes apparent and its nearby islets (Lutao or Green Island. that there are numerous disagreements Lanyu or Orchid Island, and the Penghu ar- among them. Most are also rife with no- chipelago). collecting \crtebrales and in\er- menclatural or typographical errors or cite tebrates "to provide ilic biological knowl- species names that have long since been edge required for controllini: the animal and 246 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON insect vectors of area diseases" (Secretary of the Navy Notice 5450, 9 May 1955). Over 1,900 tick collections, each containing from one to hundreds of specimens, were shipped to the laboratory of the late Harry Hoogslraal (1917-1986), then Head, Med- ical Zoology Department, U.S. Naval Med- ical Research Unit Number Three (NAM- RU-3), Cairo, Egypt, for identification and evaluation. After Hoogstraal's death, this vast resource was merged with the U.S. Na- tional Tick Collection (USNTC), now lo- cated at the Institute of Arthropodology and Parasitology, Georgia Southern University, Statesboro, while the voluminous corre- spondence pertaining to the Taiwan collec- tions was archived at a Smithsonian Insti- tution records center in Springfield, Virgin- ia. Since 1986, I have had several opportu- nities to examine critical Taiwanese speci- mens in the USNTC. With the assistance of Smithsonian archivists, I have also been able to peruse and photocopy key portions of the Hoogstraal correspondence files. The following tabulation comprises all 30 Tai- wan tick species represented in the USNTC, plus two species known from Taiwanese specimens in the Bernice R Bishop Muse- um (BPBM), Honolulu, Hawaii. Each spe- cies name appears in boldface, accompa- nied by 1 ) the total number of Taiwan-area collections of that species in, or recorded by, the USNTC or BPBM (some collections were not retained); 2) where applicable, a note on recent (and as yet controversial) classificatory changes advocated by Horak et al. (2002); and 3) a summary of all junior synonyms in earlier Taiwan checklists, or relevant remarks. I also comment on 13 tick species that, to my knowledge, do not occur in Taiwan, despite published reports or ex- tant specimens suggesting their presence. Tick Species Known to Occur in Taiwan AND Adjacent Islets Family Argasidae Argas pusillus Kohls, 1950. — 13 Taiwan collections in USNTC. Now also classified as Carios pusillus (Kohls, 1950) (Horak et al. 2002). Junior synonymy in Taiwan checklists: None. Published references to A. vesperti- lionis (Latreille, 1796) in Taiwan (Kishida 1935, Sugimoto 1939, Shimada et al. 1961, Maa and Kuo 1966, Tseng 1978) may rep- resent A. pusillus. Hoogstraal (letter no. 251, 14 February 1984) lists A. vesperti- lionis as occurring in Taiwan even though he earlier (letter no. 376, 14 February 1977) concluded that "all A. vespertilionis group samples from Taiwan . . . represent A. pus- illus." On my copy of the Hoogstraal checklist, the words ""pusillus Kohls" have been hand-entered above the typed listing for A. vespertilionis, but the latter name has not been altered. Argas robertsi Hoogstraal, Kaiser & Kohls, 1968. — 1 Taiwan collection in USNTC. Junior synonymy in Taiwan checklists: None. Published references to A. persicus (Oken, 1818) from Taiwan (Kishida 1935, Anonymous 1944, Luh and Woo 1950, Shi- mada et al. 1961, Maa and Kuo 1966, Tseng 1978) appear to be recapitulations of a mis- determined Argas. Tseng's (1978) reference to ""Argas sp. nr. arboreus Kaiser, Hoogs- traal & Kohls, 1964" (an African species) may represent the morphologically similar A. robertsi. See Hoogstraal et al. 1968a, 1975a, b. Ornithodoros capensis Neumann, 1901.— 3 Taiwan collections in USNTC. Now also classified as Carios capensis (Neumann, 1901) (Horak et al. 2002). Junior synonymy in Taiwan checklists: None. Early references to an '"Ornithodoros sp." in Taiwan (Kishida 1935, Maa and Kuo 1966) may apply to this species, but the host listed by these authors, domestic swine, would be atypical for O. capensis, which in Taiwan is known only from sea- birds. Family Ixodidae Amblyomma cordiferum Neumann, 1899. — 3 1 Taiwan collections in USNTC. VOLUME 107, NUMBER 2 247 Junior synonymy in Taiwan checklists: None. Listed only by Hoogstraal (letter no. 251, 14 February 1984). Voltzit and Keir- ans (2002) stated that this species occurs in Taiwan. Amblyomma geoetnydae (Cantor, 1847).— 3 Taiwan collections in USNTC. Junior synonymy in Taiwan checklists: Amblyomma formosanum Schulze, 1933, probably a junior synonym of A. geoemy- dae (see Camicas et al. 1998), is the name used for this species in the checklists of Schulze (1935), Elishewitz (1943), Anony- mous (1944), Luh and Woo (1950), and Shimada et al. (1961). Amblyomma helvolum Koch, 1844. — 18 Taiwan collections in USNTC. Junior synonymy in Taiwan checklists: None. Not in previous checklists, but Hoogstraal et al. (1968b) and Kolonin (1995) stated that this species occurs in Tai- wan. Amblyomma testudinarium Koch, 1844. — 46 Taiwan collections in USNTC. Junior synonymy in Taiwan checklists: Amblyomma yajimai Kishida, 1935 (Kishi- da 1935, Elishewitz 1943, Anonymous 1944, Luh and Woo 1950, Shimada et al. 1961, Maa and Kuo 1966); A. infestum tai- vanicum Schulze, 1935 (Schulze 1935, Eli- shewitz 1943, Luh and Woo 1950, Shimada et al. 1961). Note that Elishewitz (1943), Luh and Woo (1950) and Shimada et al. (1961) list both junior synonyms. As well, Elishewitz (1943), Anonymous (1944), Luh and Woo (1950), Shimada et al. (1961) and Maa and Kuo (1966) list the senior syno- nym A. testudinarium. Aponomma varanense (Supiiio, 1897).— 3 Taiwan collections in USNTC. Now also classified as Amblyomma vara- nense (Supino, 1897) (Horak et al. 2002). Junior synonymy in Taiwan checklists: None. First reported from the Penghu ar- chipelago by Robbins (1996), who de- scribed this species' convoluted taxonomic history. Elishewitz (1943) Wsts Anih/yoninui serpentinum Schulze, 1936, a junior syno- nym of Aponomma fuscolineatum (Lucas, 1847), from Taiwan, but A. varanense is the only Aponomma known from bona fide Tai- wan-area specimens. Boophilus microplus (Canestrini, 1888).— 932 Taiwan collections in USNTC. Now also classified as Rhipicephalus (Bo- ophilus) microplus (Canestrini, 1888) (Hor- ak et al. 2002). Junior synonymy in Taiwan checklists: An abundant and widespread species in Tai- wan, B. microplus has been recorded under numerous junior synonyms: B. annulatus australis (Fuller, 1899) (Elishewitz 1943. Anonymous 1944); B. annulatus caudatus (Neumann, 1897) (Kishida 1935, Ogura 1936, Elishewitz 1943, Anonymous 1944); B. australis (Fuller, 1899) (Luh and Woo 1950); B. caudatus (Neumann, 1897) (Luh and Woo 1950, Shimada et al. 1961); B. (Uroboophilus) distans Minning, 1934 (Schulze 1935, Elishewitz 1943, Anony- mous 1944, Luh and Woo 1950, Shimada et al. 1961); B. (Uroboophilus) sinensis Minning, 1934 (Schulze 1935, Elishewitz 1943, Luh and Woo 1950); Margaropus an- nulatus australis (Fuller, 1899) (Sugimoto 1939); and M. annulatus caudatus (Neu- mann, 1897) (Sugimoto 1939). Note that several checklist authors list more than one synonym. In addition, Sugimoto (1939) lists M. annulatus (Say, 1821), a junior syn- onym of B. annulatus (Say, 1821), which does not occur in Taiwan. Dermacentor taiwanensis Sugimoto, 1935._76 Taiwan collections in USNTC. Junior synonymy in Taiwan checklists: Indocentor hellulus Schulze, 1935 (Schulze 1935. Elishewit/ 1943. Anonymous 1944): Dermacentor bellulus (Schulze. 1935) (Luh and Woo 1950, Shimada et al. 1961. Maa and Kuo 1966, Tseng 1978). Haemaphysalis handicota Hoogstraal & Kohls, 1965. — 15 Taiwan collections in USNTC. Junior synonymy in Taiwan checklists: 248 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON None. Listed only by Tseng (1978) and Hoogstraal (letter no. 251, 14 February 1984). Thought to have been introduced with bandicoot rats, Baiulicota indica (Bechstein) (Rodentia: Muridae), during the Dutch occupancy of Taiwan (1624-1662) (Hoogstraal and Kohls 1965. Hoogstraal and Wassef 1973). Hae map hy salts canestrinii (Supino, 1897). — 1 1 Taiwan collections in USNTC. Junior synonymy in Taiwan checklists: None. Listed only by Hoogstraal (letter no. 251, 14 February 1984), who earlier re- corded this species from Taiwan (Hoogs- traal 1971). Haemaphysalis doenitzi Warburton & Nuttall, 1909. — 22 Taiwan collections in USNTC. Junior synonymy in Taiwan checklists: None. Not in previous checklists, but Hoogstraal and Wassef (1973) listed nu- merous collections of this species from Tai- wan and Lanyu. Haemaphysalis flava Neumann, 1897.— 4 Taiwan collections in USNTC. Junior synonymy in Taiwan checklists: None. This species appears in all previous checklists except those of Kishida (1935) and Schulze (1935). Haemaphysalis formosensis Neumann, 1913. — 27 Taiwan collections in USNTC. Junior synonymy in Taiwan checklists: None. This common species appears in all previous checklists. Recorded by Neumann (1913) from dogs in Taiwan. Haemaphysalis hystricis Supino, 1897.— 95 Taiwan collections in USNTC. Junior synonymy in Taiwan checklists: Haemaphysalis nishiyamai Sugimoto, 1935 (Sugimoto 1939, Elishewitz 1943, Anony- mous 1944, Luh and Woo 1950, Shimada et al. 1961, Maa and Kuo 1966). This spe- cies also appears as the senior synonym H. hystricis in all previous checklists, reflect- ing its prominence in the Taiwan tick fauna (Hoogstraal et al. 1965. 1973). Haemaphysalis kitaokai Hoogstraal, 1969, — 1 Taiwan collection in USNTC. Junior synonymy in Taiwan checklists: None. Listed only by Tseng (1978) and Hoogstraal (letter no. 251, 14 February 1984). The single USNTC collection (RML 16603) consists of two badly damaged fe- males obtained by Sugimoto (1937a, b) that were initially determined by Hoogstraal (1962) as H. aponommoides Warburton, 1913 (a species of the central and eastern Himalayas), but later (Hoogstraal 1969) tentatively determined to represent H. ki- taokai. The abundance of this species in southernmost Japan (Nakao and Takada 1997) also argues for its occurrence in Tai- wan. Haemaphysalis mageshimaensis Saito & Hoogstraal, 1973. — 71 Taiwan collec- tions in USNTC. Junior synonymy in Taiwan checklists: None. Listed only by Hoogstraal (letter no. 251, 14 February 1984), but recorded from numerous hosts in Taiwan, Lutao, and Lan- yu (Hoogstraal and Santana 1974). Haemaphysalis ornithophila Hoogs- traal & Kohls, 1959. — No Taiwan collec- tions in USNTC, but 8 collections in BPBM. Junior synonymy in Taiwan checklists: None. Listed only by Tseng (1978). First reported from Taiwan by Wilson (1970). Haemaphysalis phasiana Saito, Hoogs- traal & Wassef, 1974. — 16 Taiwan collec- tions in USNTC. Junior synonymy in Taiwan checklists: None. Listed only by Hoogstraal (letter no. 251, 14 February 1984). According to Saito et al. (1974), "the iaxon phasiana may pos- sibly be a subspecies of doenitzi" ', these au- thors call for additional data to clarify the status of H. phasiana in Taiwan. Haemaphysalis taiwana Sugimoto, 1936.^9 Taiwan collections in USNTC. Junior synonymy in Taiwan checklists: Haemaphysalis cornigera taiwana Sugi- moto, 1936 (Elishewitz 1943, Luh and Woo VOLUME 107, NUMBER 2 249 1950, Shimada et al. 1961, Maa and Kuo 1966, Tseng 1978). Haemaphysalis yeni Toumanoff, 1944.— 1 Taiwan collection in USNTC. Junior synonymy in Taiwan checklists: None. Not previously listed as occurring in Taiwan. The single USNTC collection (RML 77868) consists of 15 nymphs and 2 larvae removed from a specimen of the lesser coucal, Centropus bengalensis (Har- tlaub) (Cuculiformes: Cuculidae), on Lan- yu, off Taiwan's southeastern coast. The presence of H. yeni in Japan's Ryukyu Is- lands and in Vietnam (Saito and Hoogstraal 1972) suggests that it also occurs in Taiwan. Ixodes acutitarsus (Karsch, 1880). — 8 Taiwan collections in USNTC. Junior synonymy in Taiwan checklists: None. This distinctive species, said to be the largest Ixodes in the world (Yamaguti et al. 1971), appears in all previous check- lists except Ogura (1936). Ixodes granulatus Supino, 1897. — 128 Taiwan collections in USNTC. Junior synonymy in Taiwan checklists: None. Listed only by Tseng (1978) and Hoogstraal (letter no. 251, 14 February 1984). This abundant and widespread spe- cies was first reported from Taiwan by Wil- son (1970). Ixodes kuntzi Hoogstraal & Kohls, 1965. — 35 Taiwan collections in USNTC. Junior synonymy in Taiwan checklists: None. Listed by Maa and Kuo (1966), Tseng (1978), and Hoogstraal (letter no. 251, 14 February 1984). This species is en- demic to Taiwan. Ixodes nipponensis Kitaoka & Saito, 1967.— 1 Taiwan collection in USNTC. Junior synonymy in Taiwan checklists: None. Not previously listed as occurring in Taiwan. The single USNTC collection (RML 35386) consists of two females taken from domestic cattle in the vicinity of Tai- pei. Although these may have been intro- duced specimens, the presence of /. nippo- nensis in southernmost Japan (Nakao and Takada 1997) suggests that this species also occurs in Taiwan. Ixodes ovatus Neumann, 1899. — 83 Tai- wan collections in USNTC. Junior synonymy in Taiwan checklists: Ixodes shinchikuensis Sugimoto, 1937 (Eli- shewitz 1943, Anonymous 1944, Luh and Woo 1950 (as "/. shinchikuensis''), Shi- mada et al. 1961, Maa and Kuo 1966); /. taiwanensis Sugimoto, 1936 (Elishewitz 1943, Anonymous 1944, Luh and Woo 1950, Shimada et al. 1961, Maa and Kuo 1966). Note that both junior synonyms are listed in the same five checklists. The senior synonym /. ovatus is listed only by Tseng (1978) and Hoogstraal (letter no. 251. 14 February 1984). Ixodes persulcatus Schulze, 1930. — 7 Taiwan collections in USNTC. Junior synonymy in Taiwan checklists: None. Listed only by Hoogstraal (letter no. 251, 14 February 1984). Published refer- ences to /. ricinus (Linnaeus, 1758) in Tai- wan (Sugimoto 1939, Elishewitz 1943. Anonymous 1944, Luh and Woo 1950. Shi- mada et al. 1961, Maa and Kuo 1966) al- most certainly represent /. persulcatus, an eastern Palearctic sister species of the large- ly European /. ricinus. Uncommon in Tai- wan tick collections, /. persulcatus is the principal vector of Lyme borreliosis in northeastern Asia and is probably respon- sible for the small number of cases now be- ing reported in Taiwan (Shih et al. 1998, Shih and Chao 1999). Ixodes simplex Neumann, 1906. — 53 Taiwan collections in USNTC. Junior synonymy in Taiwan checklists: None. Listed only by Tseng (1978) and Hoogstraal (letter no. 251. 14 February 1984). Wilson (1970) lirsl reported /. sim- plex from Taiwan. Ixodes vespertilionis Koch, 1844. — No Taiwan collections in USNTC. but 2 col- lections in BPBM. Junior synonymy in Taiwan checklists: 250 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON None. Listed only by Tseng (1978). First reported from Taiwan by Wilson (1970). Rhipicephaliis haemaphysaloides (Su- pino, 1897). — 50 Taiwan collections in USNTC. Junior synonymy in Taiwan checklists: Rhipicephalus haemaphysaloides expeditus Neumann, 1904 (Luh and Woo 1950); R. expeditus Neumann, 1904 (Shimada et al. 1961). The senior synonym R. haemaphys- aloides is also listed by Shimada et al. (1961), as well as by Maa and Kuo (1966), Tseng (1978), and Hoogstraal (letter no. 251, 14 February 1984). Rhipicephalus sanguineus (Latreille, 1806). — 47 Taiwan collections in USNTC. Junior synonymy in Taiwan checklists: None. This ubiquitous species appears in all previous checklists. Hundreds of additional unaccessioned collections of R. sanguineus from Taiwan are in the USNTC. Excluded Species The preceding passages present argu- ments for excluding five tick species that appear in earlier checklists of the Taiwan fauna: Argas persicus, A. vespertilionis, Aponomma fuscolineatum (listed as the ju- nior synonym Amblyomma serpentinum), Boophilus annulatus (listed as the junior synonym Margaropus annulatus), and Ixo- des ricinus. Eight additional species named in these early checklists have either never occurred in Taiwan or are no longer present there: Amblyomma breviscutatum Neumann, 1899. — Listed by Elishewitz (1943) and Anonymous (1944) as A. cyprium Neu- mann, 1899. Keirans, in Voltzit and Keirans (2002), determined that the types of these species are conspecific, and since A. brev- iscutatum has page priority over A. cy- prium, the latter falls as a junior synonym. There are no Taiwan collections of A. brev- iscutatum in the USNTC. The single Tai- wanese record cited in Robinson (1926) is from "Kankan," an alternate spelling of Kaohsiung (also formerly Koshung), Tai- wan's second-largest city and its principal seaport, located in the tropical southwestern quadrant of the island. It is possible that A. breviscutatum either once occurred in Kao- hsiung (or elsewhere on Taiwan's densely populated western coastal plain) and has since been extirpated by intense urbaniza- tion, or that living specimens were sporad- ically introduced with such hosts as cattle, horses, pigs and water buffalo imported from areas to the south and west that lie well within the range of this species (e.g., the Philippines, Indonesia). Dermacentor atrosignatus Neumann, 1906.— Listed by Sugimoto (1939), EH- shewitz (1943), Anonymous (1944), and Shimada et al. (1961). In their review of this Southeast Asian species, Hoogstraal and Wassef (1985) record specimens from Palawan, Busuanga, and Culion Island in the southwestern Philippines, localities far removed from Taiwan. A single supposedly Taiwanese collection of D. atrosignatus ( 1 S, \ 2) in the USNTC (RML 15189) has been lost (L.A. Durden, letter of 28 Feb- ruary 1997 to RGR). A malformed S, also alleged to be D. atrosignatus, from "Ko- sempo" (Chia-hsien, 23.05N, 120.35E, a town in Kaohsiung County), Taiwan, was discussed by Robinson (1920). It seems un- likely that this relatively large and conspic- uously ornate Dermacentor would be over- looked in years of collecting by NAMRU- 2 investigators. Dermacentor reticulatus (Fabricius, 1794).— Listed by Maa and Kuo (1966) and by Tseng (1978). This western Palearctic species ranges no farther east than the Cen- tral Asian republics of the former Soviet Union (Pomerantsev 1950). Haemaphysalis birmaniae Supino, 1897.— Listed by Kishida (1935), Schulze (1935), Elishewitz (1943), Anonymous (1944), Luh and Woo (1950), and Shimada et al. (1961). This species, "the subject of much taxonomic confusion and many in- correct host and distribution records" (Hoogstraal 1970). is known with certainty VOLUME 107, NUMBER 2 251 only from northeastern India, Nepal and Myanmar (Burma). Haemaphy salts bispinosa Neumann, 1897. — Listed by Elishewitz (1943), Anon- ymous (1944), Luh and Woo (1950), Shi- mada et al. (1961), and Tseng (1978). This is a tropical South Asian species (Hoogs- traal and Trapido 1966); East Asian speci- mens of "//. bispinosa'' are actually H. lon- gicornis Neumann, 1901, a widespread and widely introduced tick whose original range embraced temperate areas of northeastern China, Korea, Japan, and maritime Russia (Hoogstraal et al. 1968c). Significantly, H. longicornis appears to be absent from the subtropical Ryukyu Islands (Yamaguti et al. 1971) and is therefore unlikely to occur as far south as Taiwan. Haemaphy salis inermis Birula, 1895. — Listed by Elishewitz (1943), Anonymous (1944), Shimada et al. (1961), Maa and Kuo (1966), and Tseng (1978). This is an- other western Palearctic species, ranging from southern Europe to Turkey, southern Russia, and northern Iran (Hoogstraal 1969). Haemaphysalis warburtoni Nuttall, 1912.— Listed by Kishida (1935), Schulze (1935), Sugimoto (1939), Elishewitz (1943), Luh and Woo (1950), Shimada et al. (1961), Maa and Kuo (1966), and Tseng (1978). All listings of H. warburtoni from Taiwan are apparently based on the single specimen reported by Nuttall and Warbur- ton (1915) from "Taihoku" (Taipei); this specimen was restudied by Hoogstraal (1966) and found to be H. formosensis. True H. warburtoni is restricted to moun- tainous areas in southern China, Nepal, and northern India (Hoogstraal and Kim 1985). Ixodes hexagonus Leach, 1815. — Listed by Sugimoto (1939), Elishewitz (1943), Shimada et al. (1961), Maa and Kuo (1966), and Tseng (1978). A third western Palearctic species, ranging across Europe and North Africa to western Russia and, perhaps. Central Asia (Arthur 1963). Acknowledgments For fielding innumerable questions and generously making available all Taiwanese specimens and records in their care, I thank Lance A. Durden, Department of Biology, and James E. Keirans, Institute of Arthro- podology and Parasitology, Georgia South- ern University. Thanks also to Susan W. Glenn and Paul Theerman, former Smith- sonian archivists, for providing unlimited access to the meticulously organized Hoogstraal correspondence files. For enthu- siastically encouraging the compilation of this checklist, as the first step toward real- izing Harry Hoogstraal's lifelong goal of a monograph on the ticks of Taiwan, I am grateful to Chyi-Chen Ho, Department of Applied Zoology, Taiwan Agricultural Re- search Institute, Taichung. And, as always, I deeply appreciate the assistance of Fu- Meei Yeh Robbins, who kindly prepared English translations of crucial paragraphs in Chinese papers. The opinions and asser- tions advanced herein are those of the au- thor and are not to be construed as official or reflecting the views of the U.S. Depart- ments of the Army, Navy, or Defense. Literature Cited Anonymous. 1944. Epidemiology of diseases of naval importance in Formosa. NavMed 266. Bureau of Medicine and Surgery, Navy Department. Wash- ington, DC, iv + 64 pp., appendices A-E Arthur, D. R. 1963. British Ticks. Butterworths. Lon- don, ix + 213 pp. Camicas, J.-L., J. -P. Hervy, F. Adam, and R-C. Morel. 1998. Les Tiques du Monde (Acarida. Ixodida): Nomenclature, Stades decrits, Hotes. Repartition (Especes decrites avant le 1/01/96). Editions de J'Orstom, Paris, 233 pp. Elishewitz, H. 1943. The ticks of the Pacific region. Research Project NMRI-19. Naval Medical Re- search Institute, National Naval Medical Center Bethesda, Maryland, 39 pp. Hoogstraal, H. 1962. Hcwinaphysalis ncpalciisis sp. n. from a Himalayan rodent and man, and doscri[i- tion of the male of H. n/yoiiomnioidcs Warburton (n. comb.) (Ixodoidea. Ixodidae). Journal of Par- asitology 48: 195-203. . 1966. Haemaphysalis (Allophysulis) /lospt'lo- vashtromae sp. n. from USSR and redescription of the type material of H. (A.) warburtoni Nuttall 252 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON from China (Ixodoidca, Ixodidac). Journal of Par- asitology 52: 787-K()(). . 1969. HciemaphysaUs (Alloceraea) kitaokai sp. n. of Japan, and keys to species in the struc- luraliy primitive subgenus Alloceraea Schulze of Hurasia (Ixodoidea, Ixodidae). Journal of Parasi- tology 55: 211-221. . 1970. Haemaphysalis (H.) hinnaniac Supino and H. (H.) goral sp. n. (Ixodoidea: Ixodidae), Asian parasites of artiodactyl mammals. Journal of Parasitology 56: 1227-1238. . 1971. Identity, hosts, and distribution of Hae- maphysalis (Rhipistoma) canestrinii (Supino) (res- urrected), the postulated Asian progenitor of the African leachi complex (Ixodoidea: Ixodidae). Journal of Parasitology 57: 161-172. Hoogstraal, H. and K. C. Kim. 1985. Tick and mam- mal coevolution, with emphasis on Haemaphysa- lis. pp. 505-568. In Kim. K. C, ed. Coevolution of Parasitic Arthropods and Mammals. John Wiley & Sons, New York, xiv + 800 pp. Hoogstraal, H. and G. M. Kohls. 1965. Southeast Asian Haemaphysalis ticks (Ixodoidea, Ixodidae). H. bandicota sp. n. from bandicoot rats in Taiwan, Thailand, and Burma. Journal of Parasitology 5 1 : 460-466. Hoogstraal, H. and F. J. Santana. 1 974. Haemaphysalis (Kaiseriana) mageshimaensis (Ixodoidea: Ixodi- dae): human and wild and domestic mammal hosts, and distribution in Japan, Taiwan, and Chi- na. Journal of Parasitology 60: 866-869. Hoogstraal, H. and H. Trapido. 1966. Redescription of the type materials of Haemaphysalis (Kaiseriana) bispinosa Neumann (India), H. (K.) neumanni Donitz (Japan), H. (K.) lagrangei Larrousse (Vi- etnam), and H. (K.) yeni Toumanoff (Vietnam) (Ixodoidea, Ixodidae). Journal of Parasitology 52: 1188-1198. Hoogstraal. H. and H. Y. Wassef. 1973. The Haema- physalis ticks (Ixodoidea: Ixodidae) of birds. 3. H. (Ornithophysalis) subgen. n.: Definition, species, hosts, and distribution in the Oriental. Palearctic, Malagasy, and Ethiopian Faunal Regions. Journal of Parasitology 59: 1099-1 1 17. . 1985. Dermacentor (Indocentor) atrosignatiis (Acari: Ixodoidea: Ixodidae): Hosts and distribu- tion in the Malay Peninsula, Indonesia, Borneo, and southern Philippines. Journal of Medical En- tomology 22: 644-647. Hoogstraal, H., H. Trapido, and G. M. Kohls. 1965. Studies on Southeast Asian Haemaphysalis ticks (Ixodoidea, Ixodidae). The identity, distribution, and hosts of H. (Kaiseriana) hystricis Supino. Journal of Parasitology 5 1 : 467-480. Hoogstraal, H., M. N. Kaiser, and G. M. Kohls. 1968a. The subgenus Persicargas (Ixodoidea, Argasidae, Argas). 4. Argas (P.) robertsi. new species, a par- asite of Australian fowl, and keys to Australian argasid species. Annals of the Entomological So- ciety of America 61: 535-539. Hoogstraal, H., E J. Santana, and R E D. van Peenen. 1968b. Ticks (Ixodoidea) of Mt. Sontra, Danang, Republic of Vietnam. Annals of the Entomologi- cal Society of America 61: 722-729. Hoogstraal, H., E H. S. Roberts, G. M. Kohls, and V. J. Tipton. 1968c. Review of Haemaphysalis (Kais- eriana) longicornis Neumann (resurrected) of Australia, New Zealand, New Caledonia, Fiji, Ja- pan, Korea, and northeastern China and USSR, and its parthenogenetic and bisexual populations (Ixodoidea, Ixodidae). Journal of Parasitology 54: 1197-1213. Hoogstraal. H., W. R Carney, S. Kadarsan, and R E D. van Peenen. 1973. Haemaphysalis (Kaiseriana) celebensis Hoogstraal, Trapido, and Kohls (Ixo- doidea: Ixodidae), a Wallacean member of the hystricis group: identity, distribution, hosts, and ecology. Journal of Parasitology 59: 556-562. Hoogstraal, H., M. N. Kaiser, and H. E. McClure. 1975a. The subgenus Persicargas (Ixodoidea: Ar- gasidae: Argas). 20. A. (P.) robertsi parasitizing nesting wading birds and domestic chickens in the Australian and Oriental Regions, viral infections, and host migration. Journal of Medical Entomol- ogy 11: 513-524. Hoogstraal, H., S. S. Guirgis, G. M. Khalil, and M. N. Kaiser. 1975b. The subgenus Persicargas (Ixo- doidea: Argasidae: Arga.s). 27. The life cycle of A. (P.) robertsi population samples from Taiwan, Thailand, Indonesia, Australia, and Sri Lanka. Southeast Asian Journal of Tropical Medicine and Public Health 6: 532-539. Horak, I. G., J.-L. Camicas, and J. E. Keirans. 2002. The Argasidae, Ixodidae and Nuttalliellidae (Ac- ari: Ixodida): A world list of valid tick names. Experimental and Applied Acarology 28: 27-54. Kishida, K. 1935. Notes on the Acarina-mites and ticks (including Amblyomma yajimai sp. n., on water buffalo) from the island of Formosa, col- lected in August, 1935. Lansania 7: 129-144. Kolonin. G. V. 1995. Review of the ixodid tick fauna (Acari: Ixodidae) of Vietnam. Journal of Medical Entomology 32: 276-282. Luh. R-L. and W.-C. Woo. 1950. A list of Chinese ticks. Chinese Journal of Entomology 1: 195-222. (In Chinese; English translation NAMRU-3 T19.) Maa. T C. and J. S. Kuo. 1966. Catalogue and bibli- ography of ticks and mites parasitic on vertebrates in Taiwan. Quarterly Journal of the Taiwan Mu- seum 19: 373-413. Nakao, M. and N. Takada. 1997. Survey of tick fauna in the Kyushu mainland. Japan. Medical Ento- mology and Zoology 48: 39-44. Neumann, L. G. 1913. Pediculidae, Siphonaptera, Ix- odidae. pp. 134-137. In Sauter. H. Formosa- Aus- VOLUME 107, NUMBER 2 253 beute. Entomologische Mitteilungen 2, Supple- menta Entomologica. Nuttall, G. H. E and C. Warburton. 1915. Ticks. A monograph of the Ixodoidea. Part III. The genus Haemaphysalis. Cambridge University Press, i- xiii, 349-550. Ogura, K. 1936. The ticks parasitic on the principal domestic animals in Formosa, Japan. Memoirs of the Faculty of Science and Agriculture, Taihoku Imperial University 19: 75-85. Pomerantsev, B. I. 1950. Ixodid ticks (Ixodidae). Fau- na SSSR, Paukoobraznye, new series, 4. 224 pp. (In Russian; English translation by A. Elbl, edited by G. Anastos, The American Institute of Biolog- ical Sciences, Washington, DC, 199 pp.) Robbins, R. G. 1996. Does Aponomma varanensis (Acari: Ixodida: Ixodidae) occur on the Taiwanese mainland? Journal of Parasitology 82: 672-673. Robinson, L. E. 1920. Malformations in ticks. Parasi- tology 12: 175-179. . 1926. Ticks. A monograph of the Ixodoidea. Part IV. The genus Amblyonima. Cambridge Uni- versity Press, xii + 302 pp. Saito, Y. and H. Hoogstraal. 1972. Haemaphysalis (Kaiseriana) yeni Toumanoff (Ixodoidea: Ixodi- dae): discovery in Japan, description of female and immature stages, environment, and life cycle. Journal of Parasitology 58: 950-959. Saito, Y., H. Hoogstraal, and H. Y. Wassef. 1974. The Haemaphysalis ticks (Ixodoidea: Ixodidae) of birds. 4. H. (Omithophysalis) phasiana sp. n. from Japan. Journal of Parasitology 60: 198-208. Schuize, P. 1935. Zur Zeckenfauna Formosas. Zoolo- gischer Anzeiger 112: 233-237. Shih, C.-M. and L.-L. Chao. 1998. Lyme disease in Taiwan: Primary isolation of Borrelia biirgdor- /er/-like spirochetes from rodents in the Taiwan area. American Journal of Tropical Medicine and Hygiene 59: 687-692. . 1999. Tick-borne Lyme disease in Taiwan. Chinese Journal of Entomology, Special Publica- tion 12: 161-169. Shih, C.-M., L.-R Liu, W.-C. Chung, S. J. Ong, and C.-C. Wang. 1997. Human babesiosis in Taiwan: asymptomatic infection with a Babesia microti- like organism in a Taiwanese woman. Journal of Clinical Microbiology 35: 450-454. Shih, C.-M., J.-C. Wang, L.-L. Chao, and T.-N. Wu. 1998. Lyme disease in Taiwan: First human pa- tient with characteristic erythema chronicum mig- rans skin lesion. Journal of Clinical Microbiology 36: 807-808. Shimada, T, L. W. Trager, and C. T Adams. 1961. Provisional list of the medically important fauna of Taiwan (Formosa). S"" Epidemiological FHght. Pacific Air Forces, San Francisco, California. 60 pp. Sugimoto, M. 1937a. Notes on the ticks in the For- mosan mountain-reservation for the aborigines. Journal of the Central Society for Veterinary Med- icine 50: 303-340. (In Japanese: English transla- tion NAMRU-3 Til.) . 1937b. Notes on the ticks in the Formosan mountain-reservation for the aborigines. Journal of the Taihoku Society of Agriculture and Forestry 2: 51-79. (In Japanese.) . 1939. Ticks of Taiwan. Taihoku Imperial Uni- versity, Taipei, Formosa, 63 pp. (In Japanese.) Tseng, Y.-H. 1978. Ticks of Taiwan and bordering countries. Bulletin No. 12, Ministry of Economic Affairs. Quarantine Bureau, Taipei. Taiwan. 39 pp. (In Chinese.) Voltzit, O. V. and J. E. Keirans. 2002. A review of Asian Amblyomma species (Acari, Ixodida, Ixod- idae). Acarina 10: 95-136. Wilson, N. 1970. New distributional records of ticks from Southeast Asia and the Pacific (Metastig- mata: Argasidae, Ixodidae). Oriental Insects 4: 37-46. Yamaguti, N., V. J. Tipton, H. L. Keegan, and S. To- shioka. 1971. Ticks of Japan, Korea, and the Ryu- kyu Islands. Brigham Young University Science Bulletin, Biological Series 15: i-iv, 1-226.