Ticks of Japan, Korea, and the Ryukyu Islands Noboru Yamaguti Department of Parasitology, Tokyo Women's Medical College, Tokyo, Japan

Home , Amblyomma testudinarium, Grey wagtail, Haemaphysalis concinna, Haemaphysalis hystricis, Ixodes angustus, Margaropus

Brigham Young University Science Bulletin, Biological Series

Volume 15 | Number 1 Article 1

8-1971 Ticks of Japan, Korea, and the Ryukyu Islands Noboru Yamaguti Department of Parasitology, Tokyo Women's Medical College, Tokyo, Japan

Vernon J. Tipton Department of Zoology, Brigham Young University, Provo, Utah

Hugh L. Keegan Department of Preventative Medicine, School of Medicine, University of Mississippi, Jackson, Mississippi

Seiichi Toshioka Department of Entomology, 406th Medical Laboratory, U.S. Army Medical Command, APO San Francisco, 96343, USA

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Recommended Citation Yamaguti, Noboru; Tipton, Vernon J.; Keegan, Hugh L.; and Toshioka, Seiichi (1971) "Ticks of Japan, Korea, and the Ryukyu Islands," Brigham Young University Science Bulletin, Biological Series: Vol. 15 : No. 1 , Article 1. Available at: https://scholarsarchive.byu.edu/byuscib/vol15/iss1/1

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OCT 2 9 1971 HARVARD Brigham Young University UNIVERSITY

Science Bulletin

TICKS Of JAPAN, KOREA, AND THE RYUKYU ISLANDS

Noboru Yamaguti

Vernon J. Tipton

Hugh L. Keegan

Seiichi Toshioka

BIOLOGICAL SERIES — VOLUME XV, NUMBER 1

AUGUST 1971 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN BIOLOGICAL SERIES

Editor: Stanley L. Welsh, Department of Botany, Brigham Young University, Prove, Utah

Members of the Editorial Board:

Vernon J. Tipton, Zoology Ferron L. Anderson. Zoology Joseph R. Murdock, Botany Wilmer W. Tanner, Zoology

Ex officio Members: A. Lester Allen, Dean, College of Biological and Agricultural Sciences Ernest L. Olson, Chairman, University Publications

The Brigham Young University Science Bulletin, Biological Series, publishes acceptable papers, particularly large manuscripts, on all phases of biology.

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Science Bulletin

TICKS OF JAPAN, KOREA, AND THE RYUKYU ISLANDS

Noboru Yamaguti

Vernon J. Tipton

Hugh L. Keegan

Seiichi Toshioka

BIOLOGICAL SERIES — VOLUME XV, NUMBER 1

AUGUST 1971 Financial support for Volume 15, Number 1 was furnished by the U.S. Army Medical

Research and Development Command under Grant No. DADA 1 7-71 -G-9 340. TABLE OF CONTENTS

ABSTRACT 1 INTRODUCTION 1 MATERIALS AND METHODS 2 A MAP SHOW INC GEOCRAPHICAL LOCATION OF JAPAN, KOREA, AND THE RVUKYU ISLANDS (Fig. 1) 3 KEY CHARACTERISTICS OF TICKS (Figs. 2-4) 4 ILLl'STRATED KEY TO THE GENERA OF TICKS OF JAPAN, KOREA AND THE RYUKYU ISLANDS (Fig. 5) 7 FAMILY ARC:ASIDAE 2 Ke\ to the .species of the hunily .\rgasidae 2 Genus Arg(;.« 8 Argas japoniciis Yamaguti Clifford, and Tipton, 1968 8

Argas vespcrtilionis ( Latreille, 1802) 11 GeniLs OrnithodoTOs 15 Ornithodoros capetuis Neumann, 1901 15 FAMILY IXODIDAE 18 Genius Amblyomma 18 Kev to the species of the genu.s AnMijomma 18

Amhhjommn geocmydae ( Cantor, 1847 ) 18 Amhlyommti tiitidum Hirst and Hirst. 1910 23 Ambhjoninui testudinarium C. L. Koch, 1844 25 Genus Boophilus 30

Boophihis microplus ( Canestrini. 1888 ) 30 Genui Dcrviuccntor 36 Genus Haemaphysalis 42 Kev to the species of the genus Haemaphysalts 42 Hacmaphys(dis camptinuhita \\'arburton, 1908 43 Haemaphysali-s conciniui C. L. Koch. 1844 49 Haemaphysalis sp. {H. cornigera group) 54 Haemaphyscilis doenitzi Warburton and Nuttall, 1909 59 Haemaplu/sulis fUiva Neumann, 1897 59 Hacmaphi/S(dis formoscmis Neumann, 1913 68

Haernapliysalis fujls:imi Kitaoka, 1970 68 Haemaphysalis hystricis Supino, 1897 77 Hacmaphi/salis jajioidca \Varburton, 1908 83 Hacmaptu/salis kitaokai Hoogstraal, 1969 87 Haemaphysalis longicornis Neumann, 1901 94 Haemaphysalis megaspinosa Saito. 1969 100 Haemaphysalis pental-igi Pospelova-Shtrom, 1935 105 Haem'.iphysalis wellingtoni Nuttall and Warburton. 1908 110 Genus Ixodes 112 Key to the species of the genus Ixodes 112 Ixodes acutitarsus Karsch, 1880 113 Ixodes angustus Neumann, 1899 115 Ixodes gramdalus Supino, 1897 119 Ixodes lividus C. L. Koch, 1844 124 Ixodes motws])itiosiis Saito, 1967 129 Ixodes nipjioiiensis Kitaoka and Saito, 1967 129 Ixodes oiatus Neumann, 1899 135 Ixodes persulcatus Schulze, 19.30 142 Ixodes philipi Kairans and Kohls, 1970 148 Ixodes sigtuitus Binila, 1895 150 TABLE OF CONTENTS (contimud)

Ixodes simplex simplex Neiiniaiin, 1906 155 Ixodes tanuki Saito, 1964 158

Ixodes turdiis Nakatsuji, 1942 161 | uriac White. 161 Ixodes 1852 j

Ixodes vesj>ertilionis C. L. Koch, 1844 165 I

Geniis Rhipicephalus 170 I Rhipiccpludus sanguineus Latreille. 1806 170

OOl'liTFlH. HIiCORDS 175 I

ACKNOWLEDGMENTS 177 • LITERATURE CITED 178

APPENDIX 1. NOTATIONS ON THE LITERATURE CITED 185

APPENDIX 2. COLLECTION RECORDS 186

APPENDIX 3. LIST OF MAPS AND TEXT FIGURES 219

APPENDIX 4. INFORMATION ON SPECIMENS ILLUSTRATED 222 ,

' APPENDIX 5. CORRECTED SCIENTIFIC NAMES OF HOSTS

RECORDED IN THE LITERATURE , 227

L, TICKS OF JAPAN, KOREA, AND THE RYUKYU ISLANDS

Noboru Yamaguti,' Tipton,- Vemon J. Hugh L. Keegan,'

and Seiichi Toshioka^

ABSTRACT

This publication is a revision of Ixodid Ticks rived from field collections and a thorough re- of Japan, Korea, and the Ryukiju Islands by view of the literature. Keys are provided for the

Keegan and Toshioka ( 1957 ) . The Hck fauna identification of genera and species. For each of these areas is represented by 36 species in the species we have given synonymy, a brief diag- genera Ar^fl.y, Ornithodoros, Ambhjomma, Boo- nosis, geographical distribution, hosts, and, if philus, Dermacentor, Haenmphijsails, Ixodes, known, the biology and relationship to disease. and Rhipicephalus. Three species of the family Collection data for material actually examined Argasidae and 10 species of Ixodidae were not or recorded in the literature are given in chart treated in the original edition. The additional form at the end of the paper. Illustrations of information contained in this revision was de- 65 adults and 53 immatures are provided.

INTRODUCTION

The importance of ticks as vectors of viral able. We have also chosen to present the collec- and rickettsial diseases of man has been re- tion data in chart fonn rather than as a part of viewed by H(X)gstraal (1966, 1967a, 1967b). the text. The format used by Elbl and Anastos

Ticks are also important vectors of pathogens ( 1966) has been used as a guide in the prepara- affecting domestic and game animals. The man- tion of the collection data chart. ual, Ixodid Ticks of Japan, Korea, and the Rt/u- Much of the infoniiation contained in this ktju Islands by Keegan and Toshioka ( 1957), rep- paper has been copied verbatim from the origi- resented a significant contribution to our knowl- nal 1957 manual. The order in which the edge of tick vectors of disease. Inasmuch as authors are listed does not infer that Keegan and there have been several biological and taxonomic Toshioka have made any less contribution than studies of the tick fauna of Japan published since the other two authors, and as a matter of fact 1957, and to a lesser extent Korea and the this paper would hardly have been possible Ryukyu Islands, Colonel Keegan suggested that without the original manual as a starting point. Tipton and Yamaguti revise the original paper. Furthennore, we received constant encourage- We have supplemented information from the ment and assistance from Colonel Keegan and literature with our own collecting and life cycle Dr. Toshioka. studies at the 406th Medical Laboratory. Some As in the 1957 manual, the generic classifica- additions and changes have been made in the tion given by Anastos (1950) and Hoogstraal format, such as the inclusion of the argasid ticks, (1956) has been followed. We have also relied a diagnosis for each species, an illustrated key to heavily on the assistance of Mr. Glen M. Kohls the genera, and a map for each species showing ( Rockv Mountain Laboratory, Hamilton, Mon- distribution based on all collection records avail- tana), Dr. Harry Hoogstraal (Naval Medical

*[)eparlnient of Parasitology. Tokyo Women's Medicil (College. Tokyo. Japan. -Departnient nf 7.noIogy. Rrigliani YoiinR Universily. I'rovo. Utah. 'ncpartnifnt of Prpveiitive Mi'dii inn. Sdionl of MedMine. University of MississiiJpi. Jatksoii. Mississippi. 'Department of Kntomologw -HHuli .MeJital I.aboratory. U.S. Army ,\Iedical Command. .\P0 San Francisco. 06543, US.\. Bhigham Young University Science Bulletin

Research Unit No. 3, Cairo, Egypt), and Dr. MATERIALS AND METHODS Shigeo Kitaoka (National Institute of Animal Most of the material examined as a part of Health, Tokyo). this study was collected during extensive field Taxonomieally important striictnre.s are trips on the four main islands of Japan and on labeled in Fig. 2 to 4. A detailed discussion of islands of the Ryukyu Archipelago. A few speci- tick morphology is not given in this paper. Such mens were provided by one of us (Tipton) in infomiation may be obtained from texts of medi- connection with field excursions made to Korea. cal entomology or the excellent monographs of Engorged females were kept in vials until Nuttall and Warburton (1908, 1911, 1915). oviposition, and larvae were fed on ears of rab- Synonymy listed under the specific name in- bits so that laboratory life cycle studies could be volves only those papers in which specimens initiated. Unengorged females ( freshly collected from Japan, Korea, or the Ryukyu Islands were from the field) were likewise reared in the examined or discussed by those authors. Though laboratory. These procedures enabled us to there have been several papers dealing with properly associate adults with immature forms. specimens from Taiwan ( Formosa, previously Tentative identifications were recorded on cards controlled by Japan before World War II), they with collection data. One card per host was were purposely excluded as sources of synonyms prepared. Specimens from each lot were sent and literature, except in those cases where data to Mr. Glen M. Kohls, Rocky Mountain Labora- contained were pertinent to the populations in tory, for confinnation of identifications. the areas discussed. Hence, the synonymy pre- Unmounted specimens collected in Japan, sented here is less extensive than that of other Korea, and the Ryukyu Islands and preserved monographic papers. Literature containing bio- in 70? alcohol were used as a basis for most of logical information germane to this paper is the illustrations. Often it was necessary to re- included. move legs and spiracular plates in order to get a clear view of contour and chaetotaxy. In With practicality in mind a brief diagnosis of some instances the capitulum of the adult and each species is given rather than a detailed whole body of the nymph or larva were mount- description. Keys to the immature forms are not ed temporarily in Hoyer's solution on a cavity included, but many illustrations of larvae and slide for detailed examination of the chaetotaxy nymphs, as well as adults, which were not pro- and dentition of the hypostome. vided in the publication of Keegan and Toshioka Illustrations were prepared by the artists (1957) have been added. We believe this will named in the acknowledgments. be beneficial to field workers concerned with tick biology. Family Argasidae Canestrini, 1890

At the time of the writing of this paper Nonscutate "soft ticks." Sexual dimorphism (1970), the political boundary bet^veen Japa- very slight. Integument of adults and nymphs nese and American administration of the Ryukyu mammilated, wrinkled, leathery, granulated, or Islands lies between Amami Gunto and Okinawa with tubercles. Capitulum of adults and nymphs Gunto, the former having already reverted to in camerostome on ventral side; apart from Japan. In 1972, the remaining islands in Oki- anterior margin of body. Palpal articles sub- nawa and Sakishima Gunto ( including Miyako equal, leglike, free, never fused in all stages. and Yaeyama Retto) will return to Japanese Porous areas absent. Eyes present or absent; administration, probably as Okinawa Prefectiire. when present, on the supracoxal folds. Spiracles Even though Amami Oshima is politically Japa- of adults and nymphs usually anterior to coxae nese, for the purpose of geographical clarity, it IV. Pulvilli usually absent or nidimentary in is shown in Appendix 2 as being part of the adults and nymphs, whereas functional in larvae.

Ryukyu Islands. It is administered by Kago- Number of nymphal stages variable. Type shima Prefecture in Kyushu. genus: Argas Latreille, 1796.

Key to the Species of the Family Argasidae

1. Suture between dorsal and ventral surface of body absent; dorsal surface mammilated;

on seabirds (Fig. 9, 10) . Ornithodoros capensis Suture between dorsal and ventral surface of body definitely present; dorsal surface finely wrinkled Genus Argas 2

2. R()d\ iicarK- round; on bats (Fig. 7. 8) Argas vesjwrtiUonis Body longer than wide; on swallows (Fig. 5, 6) A. japonicus Biological Slhies, Vol. 15, No. 1 Ticks

Maf I. Map of Japan. Kore;i and the Rynkyii LslaiuLs. Bricham Young University- Science Bulletin

capi tulum

cervical pit cervical groove

punctation pregenital plate

scutum - genital aperture

lateral groove - median plate

epimeral plate

adanal plate

anal plate

hypostome

porous area internal spur

scapula coxa I

lateral carina coxa II

- cervical groove - coxa III

scutum -coxa IV -external spur marginal groove spiracular plate "genital aperture genital groove -anus

anal groove

Dorsal view of female Ventral view of female

Fig. 1. Key characteristics of ticks - 1 (genus Ixodes) Bi()i,oc:iCAL Series, Vol. 15, No. 1 TicK.s

internal spur

external spur eye ornate marking

fovea

accesory shield festoon

adanal shield

postanal -median groove

Female of genus Amblyomnia (dorsal view) Male of genus Rhipicephalus (ventral view)

hood

camerostome cheek — eye capitulum genital aperture eye

suprocoxal fold

spiracle

anus

preanal groove

transverse postanal groove

genus Ornithodoros

Fig. 2. Key characteristics of ticks - 2. i

Bhigham Younc University Science Bulletin

article III -external profile

article II __ postero-external juncture

-- article I --dorsointernal setae

basis capituli

--porous area-

cornua

-hypos tome

article IV

infrainternal or ventrointernal setae

-article I -auricula

basis capi tul genus Ixodes genus Haemaphysal is

pul villus claw hump apicoventral spur macula or hook Mailer's organ

- goblet

posthypostomal seta

Hypostome Tarsus I Spiracular plate

Fig. 3. Key characteristics of ticks - 3. k

BioLotiiCAL Series. \*oi,. 15. No. 1 Ticks

Famil y Ixodidoe Famil y Arqosidoe Capitulum at anterior end of body. Scutum or dorsal shield present.

Capitulum on under side of (Short in? Long in «f ) body. Scutum absent.

Usually with definite sutural line Sutural line absent. Body separating dorsal and ventral sur- margin structurally not faces; body margin distinctly different from remainder flattened and structurally differ- of integument. ent from remainder of integument.

/ Anal groove contouring the anus Anal groove contouring the anus posteriorly, festoons present. anteriorly, festoons absent.

Argos Ornithodoros

Eyes absent. Scutum inornate. Second Eyes present. Second segment segment of Palpi usually projecting of Palpi not projecting later- laterally beyond the basis capituli ally beyond the basis capituli. except H. itaokai ?

Palpi long and slender, longer than Palpi short and broad, not longer Haema ph ysalis the basis capituli. Scutum ornate. than the basis capituli.

Basis capituli rectangular. Scutum Basis capituli hexagonal ornate. Coxa IV of male much larger than other coxae. Amblyommo

I Palpi very short Dermacentor and ridged dorsally Palpi not unusually short, not ridged. and laterally. Coxa I with two very Coxa I with two long spurs. Festoons short spurs. Festoons absent. present.

Coxa I Coxa I

? J tL kitaokai

BoQphilus

Fio. 4. Illustrated key to the genera of ticks of Japan, Korea, and the Ryiikyu Islands. )

BnioHAM YouNO University Science Bulletin

Genus Argas Latreille, 1796 A. vulgaris Filippova, 1961, of eastern Russia, but adults may be differentiated from these Body distinctly flattened dorsoventrally; dor- species on the basis of number and distribution sal and ventral surfaces subequal in area. Body of setae on the tarsi, basis capituli, and anal margin flattened even when depleted, consisting valves. Larvae may be diftcrentiated on the of radial striations or <|uadrangular plates. Su- basis of length, number, distribution, and stnic- tural line separating dorsal and ventral surface ture of body setae (A. lagenoplastis); dentition present. Integument leather)' or finely wrinkled, of the hypostome and number and distribution intermingled with small "buttons"; discs ar- of body setae on the posterior (juadrant (A. ranged more or less radiallv. Eyes absent. cooleiji; and dentition of the hypostome and Adults and nymphs similar. Parasitic principally size of the dorsal plate (A. vulgaris). on birds or l)ats. Type species: Argas reflexiis This swallow argasid is easily distinguished (Fabricius, 1794). from other soft ticks of the Japanese fauna in Argas japonictis Yamagnti, Clifford, and Tipton that the body is ovate, the ventral "paired organs" are lacking, the mouth parts are not so (Fig. 5 and 6) close to the anterior margin of the body as in Argas japoniciis Yamaguti, Clifford, and Tipton, vespertilionis, and the tarsi have distinct sub- 1968:453-459, Fig. 1-17; Uchikawa and Sato, apical dorsal protuberances. 1968:157-161, Fig. 1, 1969:95-97; Saito, lijima, and Minai, 1969-39-41. DlSTRIBUTIGN AND HoSTS: Argas .sp.: Inatomi and Yamaguti, 1960:17-18; To date this species is known only in Japan Uchikawa, Sato, and Kugimoto, 1967:141- and Korea. Reported hosts are the Japanese 151, Fig. 1-20. striated swallow, Hirundo daurica japonica. and Argas var. japonicus reflexus Yamaguti and Ina- the Japanese house martin, Deliclion urbica tomi, 1961:142. da.njpus. Under experimental conditions A. Argas persicus (not Oken, 1818): Hara, 1963: japonicus will feed on chickens. 123-125, Fig. 1.

Discussion: ArgdS japonicus

The occurrence of this soft tick was first re- 9 Collected and examined ported by Inatomi and Yamaguti ( 1960 ) from nests of the Japanese striated swallow, Hirundo O Collection records from literature dattrica juponica, which were built under eaves 3 Combination of above ft of the post office at Niimi City, Okayama Pre- O fecture, Honshu. They gave a brief account of adult morphology. The following year, Yama-

guti and Inatomi ( 1961 ) examined immature forms from this locality and considered this population to be a new variety of Argas reflexus, which they designated japonicus.

Hara ( 1963) reported the occurrence of Argas persicus from nests of the house martin at Agematsu, Nagano Prefecture. Uchikawa, Sato,

and Kugimoto ( 1967 ) found a large number of specimens of Argas sp. in swallow nests built under eaves of the University Hospital of Shin- shu University, Matsuinoto City, Nagano Pre- fecture. Yamaguti, Clifford, and Tipton ( 1968 examined this material, including Hara's persicus. and indicated that all specimens were identical to the new species, A. japonicus, described by them.

Diagnosis:

This species is similar to A. coolet/i Kohls and Hoogstraal, 1960, of western North America, A. lagenoplastis PVoggatt, 1906, of Australia, and Map 2. Known distribution of Argas jtiponUus. Biological Series, Vol. 15, No. 1 Ticks

X A^niia.

Fic. 5. Argas japonicus, female and male. Bhicham Young University Science Bulletin 10

-r A-^Ac

Fig. 6. Argas japnnicus, lana ani> nvTnph. Biological Series, Vol. 15, No. 1 Ticks H

Biology: Disease Relationship:

Uchikavva et al. (1967) and Uchikawa and There are .some reports of invasion of dwell- Sato (1968, 1969) examined a natural popula- ings by this tick, and inhabitants have often tion in Nagano Prefecture and studied a labora- complained of tick bites (Inatomi and Yama- tor\' colony led on chicken.s. Tiiey found the guti, I960; Hara, 1963; Uchikawa et al., 1967; tick.s to be very active, and .some of them in- and Uchikawa and Sato, 1969). However, there truded into the wards of the hospital (Shinshu is no definite evidence of biting by A. japonicus, UniversitA' Hospital) during the period from late and it may be that bites were caused by the March to September when the host birds were fleas, FwntopstjUa setiger Smith or Ceratoplnjl- nesting. .Ml de\eloping stages, as well as eggs, Itis farreni chaoi Smit and Allan, which were were found in the holes and crevices of mud often associated with A. jajwnicus in swallow used in construction of nests. After the host nests. birds leave and migrate to southern regions, The relationship of this tick to human dis- the ticks stav in the same niches and overwinter ease is not known, but because it lives in close there. Uchikawa et al. (1967) state that the proximity to man the potential for transmission winter colonv consists of unfed specimens plus of disease to humans is real. those which have fully digested a previous blood In 1960 Yamaguti found a nestling bird meal. Unfed larvae die within three weeks heavily infested with larvae and nymphs of A. when kept at 30 C, but they survive until the japonicus. The bird appeared to be weakened following spring without a blood meal under to the point of death. natural conditions. There are at least two instars Argas vespertilionis (Latreille) in the nvmphal stage, and both are often found (Fig. 7 and 8) in natural populations. In laboratory colonies 4th instar nymphs are seen only occasionally. Caris vespertilionis Latreille, 1802/1803:67-68; Morphological differences among nymphs of dif- Kishida, 1936:142. ferent stages are generally very slight. Uchi- Argas vespertilionis: Kishida, 1927:986, Fig. 1899, kawa and Sato ( 1968 ) studied the morphological 1947:975, Fig. 2775; Saito, 1955:7-12, Fig. differences of these nymphs and found that the 7-14; Kamo, 1962:173; Kishida and Asa- apicoventral setae on the tarsi change with numa, 1965:395, Fig. 214; Asanuma, 1965a: molting, and this serves to discriminate the 124; Yamaguti, Clifford, and Tipton, 1968: nymphal instars. In the laboratory Uchikawa 453. et al. (1967) found that larvae, nymphs, and Argas sp. Ito and Saito, 1954:563-564; Saito, adults began to feed on chickens after a short 1955:7-12, PI. II. preparasitic period of about 3 days. Larvae fed for 3-6 davs, and nvmphs and adults for 9 min- Discussion: utes to an hour. At 30 C the postparasitic period The first record of A. vespertilionis in Japan for larvae was 5-12 days, for 1st instar nymphs is probably Kishida's account in the 1927 edition it was 8-18 davs, and 2nd and 3rd instar nymphs of the Illustrated Encyclopedia of the Fauna which molted into adults it was 12-20 days. The of ]apan. He stated that this species occurs in period was prolonged at lower temperatures. Honshu, Shikoku, Kyushu, Korea, and the Ryu- Each female laid .36-200 eggs during a 5-12 day kyu Islands and listed seven species of bats as period, and the larvae hatched in 12-16 days. host animal in the above areas. Kishida (1936) At the 406th Medical Laboratory rearing ex- also reported his collection of Caris vespertilionis periments on chickens were undertaken under in Korea. Saito ( 1955 ) collected Argas sp. from conditions somewhat different than those de- bats in Niigata City and reared the species on scribed by Uchikawa et al. (1967). Newly the host, Pipistrelhis ahramus. He did not defi- liatched larvae were reared individually, and nitely identify the tick, but he mentioned that his some of them emerged to adults after passing material was very close to A. vespertilionis. Al- through the 4th nvmphal instar. Most of the though specimens were not available for study, larvae developed to adults, but they remained morphological characters shown in photographs in a postparasitic (|uiescent phase without ovi- of the tick are similar to those of vespertilionis. positing for more than one year. After feeding, Additional material h;is been collected from the bodv outline of the larvae changed and Pipistrelhis spp. at Fukuota, Kyushu, and Sap- closelv approached the nvmphal shape. The poro, Hokkaido, and from Vespertilio superans details of rearing experiments will be sum- at Shojo, Fukushima Prefecture, Honshu. Tlie marized and reported elsewhere in the near lot from Ves])ertiUo superans was sent to Kohls future. and it was his opinion that the specimens differ Bricham Young University Science Bulletin 12

Fic. 7. Argas vespertilionii, female and male. 13 Ticks Biological Series. Vol. 15, No. 1

Fig. 8. Argas vespertilionis, lana. 14 Bhigham Young University Science Bulletin only slightly in integumental characters from Egyptian vespertilionis and may well be the same species.

Hoogstraal ( 1956) considers the A. vespertilionis group to include European and African vesj)ertilionis, A. pussihis Kohls, 1950 on Palwan Island in the Philippines, and closely related populations of uncertain species status, and that this group ranges tliroughout the continent and island groups of the world except in the Americas.

Diagnosis:

This round bat-tick is the sole argasid species parasitizing bats in Japan and Korea and is readily distinguished from other argasids in this fauna. The Ixxly is circular, the anterior margin has a very slight anterior projection, in the peripheral area there are rectangular "cells" and a definite sutural line dividing the dorsal and ventral surfaces, there are distinct ventral "paired organs" posterior to the anus, and the mouth parts are close to the anterior margin of the body.

Distribution and Hosts: Hoogstraal (19.56) gave the distribution and hosts of the vespertilionis group as England, the Netherlands, Sweden, Spain, Germany, Africa, Korea, China, the Philippines, and Ceylon. The Maf .3. Known distribution of Ar<^iis lespcrtilionis. group is also known to occur in southern India, Cambodia, Australia, France, Italy, Russia, and after the second meal. Nymphs become re- Japan. Sugimoto (19.36b) cited Sauter's collec- plete in 20-50 minutes, usually .30-40 min- tion of A. vespertilionis on Taiwan. Almost all utes. Adult males may develop from the species of bats within the distributional range first nvmphal molt, but usually nymphs molt of A. vespertilionis may be parasitized by this twice before becoming adults. Males and tick, and all stages of the tick mav infest these females may feed within 7 davs after molt- bats. Both nyiuphs and adults occasionally at- ing, and they feed for .30-40 minutes. No tack man. females oviposited within 6 months after the nvmphal-adult molt, even though with Biology: males continuously and given two to six Saito (1955) reared this species on the host blood meals. The first oviposition follows a bat in the laboatory but did not mention details blood meal bv about a week and appears of the life cvcle, except that the nvmphs fed at to trigger a physiological release mecha- night, adults appeared about three months after nism, because in several instances females passing through three nymphal instars at 25-.30 have deposited fertile eggs three months C, and that below 10 C the ticks of each stage afterwards with or without a meal. went into hibernation. Hoogstraal (1956) reported that A. vesper- Disease Relationshii': tilionis was succ-essfully reared in his laboratory Nvmphs and adults may attack man (several at 80-90 F and 40-50? relative humidity. authors), and in Japan, Kamo (1962) reported Eggs hatched 16-20 days after oviposi- one instance of a human lieing bitten by this tion and larvae fed as soon as 4 days after- tick. Reports indicate that mild itching may wards. Larvae fed for 14-31 days, usually persist for several weeks. This tick may be a 17-19 davs, and molted 5-10 davs later. vector of a spirochete of bats, but conclusive Nvmphs fed 3-4 davs alter molting and they supporting evidence is not available. The rela- usuallv fed twice, followed bv a molt S or tionship of this tick to human disease has not 9 days after the first meal and 12-14 days been studied. Biological Sebies, Vol. 15, No. 1 Ticks 15

Genus Ornitlioduros C. L. Kocli, 1S44 known to occur. These include islands off the coasts of South Africa, southern Australia and Body more or less flattened. Dorsal surface Japan, and additional islands in the Atlantic, usually con\'ex when fully fed. Body margin Pacific, and Indian oceans. structurally not different from adjacent areas. Amerson (1968) states: "It is presently Suhiral line absent. Capituluni suhtenninal or known from 32 islands and at sea in the Central apart from anterior margin of the body. Hood, Pacific and immediate surrounding areas from cheeks, and camerostoine present. Dorsal humps 22 sea and shore bird species, as well as from and protuberances prominent on tarsi. Eyes man, the European rabbit, sea turtles (Celonia absent or present.

mijdas), and ground litter . . . Since 1901 O. Type Species: Oniithodoros s(ivi<;^mii (Audouin, capensis has been recorded aroimd the world 1826). in the tropical and temperate regions and has been associated with 29 species of sea and shore Ornithodows capensis Neumann birds throughout the world (from published (Fig. 9 and 10) and unpublished records)." Amerson also gives Ornithodows talaje var. capensis Neumann, a world distribution map of this species. 1901:2.58. In Japan O. capensis has been found on the Ornithodows cajyensis Neumann, 1901; Kohls, black-tailed gull. Lams crassirostris, the streaked 1957a: 89-90, Fig. 1; Keegan and Toshioka, shearwater, Calonectris Jeucomelas. and the an- 1957:23; Asanunia, 1960:94, 1965a: 124, Fig. cient auk, Synthliborainplnis antiqitus. Accord- 7.16. ing to Asanuma (1960), O. capensis will attack domestic fowls. DiSClTSSION: The original description of this species was Biology: based on material taken from nests of penguins O. capensis has been reared on chickens at on islands off the coast of Cape Colony, South the 460th Medical Laboratory. Larvae fed for .\frica. Kohls (1957a) considers capensis to be 5-6 days and molted to 1st instar nymphs 1-4 distinct enough to warrant full specific status and to be readily distinguishable from talaje on the basis of morphological differences as well as host relationship and distribution. In Japan, O. cai>ensi.^ was first found by Asanuma associated with Ixodes signotus in July 1955 at Kabujima, Aomori Prefecture, Honshu. He did not, however, refer to this soft tick in his two papers on Ixodes signatus published in 1955 and 1957. In his paper of 1960 Asanuma assembled the known information on O. ca})ensis in Japan up to that time and indicated he had collected this species in 1955. Later the tick was collected by 406th Medical Laboratory personnel at the same lo- calitv', as well as from several other localities.

Dl\gnosis:

This soft tick is the only species of the genus OrnitJiodoros parasitizing sea birds in this area and is easily recognized by the generic characters used in the key. As reported by Kohls (1957a), this species resembles O. amhhis, but differs from that species in having a more pointed anterior margin of the body and cheeks with partially overlapping mouth parts.

Distribution .\nd Ho.sts:

O. caj)ensis is known to be widely distributed

throughout the world. Kohls ( 1957a ) reviewed the distributional records of this species and listed the names of islands where the species is .M.-^p 4. Known distribution of Ornithodoros capensis. BniGHAM Young Univehsity Science Bulletin 16

2.0 mm

0.5 mm 0, 5 mm

0.5mm

Fig. 9. OrnitJwdoros capensis, female and male. Ticks 17 Biological Series, Vol. 15, No. 1

0.02 mm

^i^>>^'//^^y^y^^^

- A^Wo

Fig. 10. Ornithodoros capensis, larva and nymph. 18 Brigham Young University Science Bulletin days afterwards. Ist instar nymphs molted to (behind capituluin) in females, nymphs, and 2nd instar nymphs in 13-17 days without a blood larvae. C-'apituhim at anterior margin of body. meal. These 2nd instar nymphs, when given a Porous areas present in female on dorsal side of blood meal within 9 days after molting, molted basis capituli. Eyes present or absent; when again to 3rd instar nymphs 14-18 days after the present, laterally on the scutum. Palpal articles blood meal. There were at least three instars in not free; 4th article much reduced, inserted ven- the nyinphal stage under laboratory conditions trallv on 3rd ;irticle. Spiracular plate present, at approximately 25 C, but some 3rd instar posterior to coxae IV. nymphs developed into 4th instar nymphs. De- „ „ i / i i_ n i-rnc -., ' ' Type Genus: /xor/e.s Latreillc, 1795. r 1 IT I 1 11 1 .11 I, tails of the life cvcle shall be reported elsewhere at the conclusion of current rearing and life- Genus Ambhjomina C. L. Koch, 1844

' ^ Generally ornate. Palpi long; article II at Disease Relationship: least twice as long as wide. Eyes and festoons Unknown. present. Basis capituli variable in form; usually roughly triangular or rectangular dorsally. Family Ixodidae Murray, 1877 Adanal shield absent in male. Spiracular plate

. rousihlv triangular or comma-shaped. ,.,,.,„ ^ , ,. I o i Scutate hard ticks. Sexual dimorphism ^ . marked; well-sclerotized scutum covering the Type Species: Amhhjomma cajennense (Fabri- dorsal surface almost entirely in males, partially cius, 1787).

Key to the Males of the Genus Amhhjoinina

1. Hypostome with denticles of inner file almost equal with others; coxae II, III, IV with subecjual external .spurs; on tortoises (Fig. 12) geoeimjdae Hypostome with denticles of inner file much smaller than others; coxa IV with external spur much longer than those of coxae II and III; on larger wild and domestic mam-

mals ( Fig. 17) testudinarium

Key to the Females of the Genus Amhhjomma

1. Hypostome with denticles of inner file much smaller than others; coxa IV with external spur slightly longer than those of coxae II and III; on larger wild and domestic mammals (Fig. 16) testudinarium Hypostome with denticles of inner file almost equal with others; coxae II, III, IV with subecjual external spurs 2

2. Coxa I with two spurs; body with thick setae; dorsal foveae unusually large; on tortoises (Fig. 11) geoemijdae

Coxa I with single small spur; body without thick setae; dorsal foveae inconspicuous; on .sea snakes (Fig. 15) nitidum

Amhhjomma geoemijdae (Cantor) and noted morphological characters, but he did (Fig. 11-14) not mention the relationship betvveen the two, of "t'OCHii/f/we were , , , ,, ,„,~ r.^r, probably because specimens Ixodes oeoemyduci..mor. 1847:608. \^^^ ,^^^,(.^^^^^ ^^^ l^^^ Later Kohls (1957b) Amhhjomma maknjunum Neumann, 1908:14-16, discussed both species 'and showed mahnjamim Fig. 9-10; Toshioka, 26, Keegan and 1957:8, (^ ^e a svnonvm of oeocmiidac. PI. Asanuma, 4, 5; Kawashima, 1963:103; Specimens reported here were ail taken from 107. 1965a: tortoises on Kyushu and in the Hyukyu Islands. D,o^.,oo,^», One specimen, a nvmph found on a human, was iscussiON: 1,1 r 1 .1- stray a tortoise examined the . probably . from „, , 1, , 1 , 1 ^, a '. I his tick, originally described by (>antor m ^ i/ IS47, was collected from the neck of Cxeocmifda ' '^' spinosa in Sumatra. Keegan and Toshioka Diagnosis: (1957) collected males, females, and nymphs This tortoise tick is easily distinguished from from tortoises on Kyushu and in the Ryiikyu nitidum and testudinarium by the combination Islands. Anastos (1950) reviewed the synonymy of characters used in the key. The female of and distribution of mahnianum and '^eoemrjdae this species is especially distinctive in that the Biological Series, Vol. 15, No. 1 Ticks 19

Fig. 11. Amblyornma geoemydae, female. University Science Bulletin 20 Bhigham Young

Fig. 12. Ambh/omina geoemijdcw, inal Biological Series, \'ol. 1.5, No. 1 Ticks 21

Tti. TrLc^^iu,

Fig. 1.3. Amblyomma geocmt/dae, nymph. Brigham Young University Science Bulletin

'?n. Tm^-tiA^ 0.2 mm

Fic. 14. Aml>li/omma gcocvujdae , larva. BioLor.icAi, Series, Vol. 15, No. 1 Ticks 23 clor.suin hii.s a pair of large, adjacent foveae on the post.scutal area; the dorsal surface has thick, short setae; and the basis capituh has large porous areas.

DisTiuBUTiox .\Ni) Hosts: Anastos (1950) stated that the species was known only from tortoises collected in Singa- pore, Malaya, Simiatra, and the Philippines. Kohls (1957b) also reported this species from Sarawak ex Varanus salvator, Ctjlemtjs dentata (as C. (Ihor), Ueosemijs spinosa (as Geoemijda spinosa), Te.studo cmtjs, and "stomach of fish," Pontius sealei.

Amblyomma geoemydae . ," ! Bricham Young University Science Bulletin 24

Fig. 15. Amhhiommn nitUhim. female. Biological Series, Vol. 15, No. 1 Ticks 25 not have marginal groo\'e.s; and the dorsal foveae Disease Relationship: are inconspicuou.s. Unknown. Distribution and Hosts: Amblyomma testudinarium C. L. Koch The knowai distribution includes: Solomon (Fig. 16-19) Islands, ex sea snake (Hirst and Hirst, 1910); Amhlijomma testudinarium C. L. Koch, 1844: Andaman Islands, host unmentioned (Sharif, Kishida, 1922a: 850-851, Robinson, 1928); Singapore ex Laticauda coluhrina (War- 226; 856; 1926:257; Sugimoto, 1937a: 317-323, 1937b: burton, 1932, as A. laticimdae: Audy et al., 612-613, PI. 4; Nakamura and Yajima, 1937: 1960); and the first record from the Ryukyu Fig. 1-9 in PI. Itagaki, Noda, Islands reported herein. 174-175, \IV; and Yamaguchi, 1944:1-97, PI. 17, 22, 1959: 1-118, PI. 17, 22; Keegan and Toshioka, 1957:

9, PL 6, 7; Kawashima, Kamo, and Miyazaki, 1960:77-80; Tanaka et al., 1960:69-70, Fig. 1-3; Nagahana and Matsuo, 1962:119-120; Asanuma, 1965a: 106-107, Fig. 7.3, 1965b:400, Fig. 226. Ambhjomma ijajimai Kishida, 1935:137-138, Fig. 1-4; Nakamura and Yajima, 1937:176, Fig. 7, 8 in PI. I, Fig. 5-10 in PI. XV; Itagaki, Noda, and Yamaguchi, 1944:1-97, PI. 18, 22, 1959:1-118, PI. 18,22.

Discussion: Anastos (1950) and Kohls (1950) have reviewed the extensive synonymy of this widely distributed species. The first report of this tick from Japan (Kishida, 1922a) was based on a specimen collected from a pig in Tokyo. Sugi-

moto ( 1937a, 19.37b ) reported two collections of larvae from the frog, Microhtjh fissipes, and from a green frog (sp?) on Ishigaki, an island in the Ryukyu archipelago. These two lots were not collected by Sugimoto but submitted to him by the collector. Considering the host-parasite relationship of this tick, it is very questionable that A. testudinarium was parasitic on a frog. In his description of Ambhjomma ijajimai Kishida (19.35) included illustrations of venter, scutum, spiracular plate, and the 1st tarsus of M.\p 6. Known distribution of Ambhjonima nitiduv the holotype female. He gave only diagnostic

characters to separate it from A. testudinarium, Biology: and a precise description of the holotype was It is of interest that this nocturnal sea given later by Nakamura and Yajima (1937). snake spends the daytime resting, often in The holotype specimen labeled as "1291 l.VII. tightly-packed groups, in mangrove tree- 1935. Taito-cho, Hinan-ku (Formosa), Pasicau, holes or in rock crevices out of the water. from mammae of water buffalo, Yajima Am- Furthemiore, the genus Laticauda appears hh/omma ijajimai Kishida" was examined by

to have evolved from an elapid stem much Kohls, and it was his opinion that A. ijajimai is

later than the other sea snakes. This would is conspecific with A. testudinarium Koch. A.

explain not onlv the infestation by Am- Ijajimai was also reported by Itagaki et al. ( 1944, hh/onuna, but also the infestation of the air- 1959), and all of these collections are from water sacs by larvae of a peculiar trombiculid buffalo, Buhalus hubalis, on Formosa. mite, Vatacarus ipoides Southcott ... It will be interesting to discover how A. nitidum Diagnosis:

is adapted to parasitizing this sea snake This large tick is easily distinguished from (Audyetal., I960). nitidum and geoemijdae in that the denticles of Brigham Young University Science Bulletin 26

Fig. 16. Amhli/omma testudmiirium. female. BrOLOGiCAL Series, \'ol. 15, No. 1 Ticks 27

Fic. 17. Amhlyommii testudinarium, male. 28 Brkjham Youno University Science Bulletin

^^r-

'M.. ^ti»»^

Fic. 18. Atnhlyomma testudinarium . nymph. Bk)i,oc.icai. Sehies, Vol. 15, No. 1 Ticks 29

KiG. 19. Amhlyoinrna tcstudinarium, larva. 30 Bhigham Young University Science Bulletin

the inner file are much smaller than those of the a wide variety of wild as well as domestic ani- other files; the external spur of coxa I is longer mals. One of the unengorged females collected than the internal spur; the spur of coxa IV is by 4()6th laboratory personnel from Amami longer than those of coxa II and III which arc Oshima was placed on a rabbit's ear in the short and blunt in the male, broad and ridgelikc hiboratory on 6 November 1967, fed for a week, in the female; tarsi III and IV have two distinct and increased in size to 23 mm in length and subapical ventral spurs; the dorsum has a pair 21 mm in width. After engorgement this tick of small foveae which arc well separated in the died on 26 November 1967 without laying eggs. female. Disease Relationship: Distribution and Hosts: "Krijgsman and Ponto (1932) stated that

Kohls ( 1957b) has listed the distribution and this species transmits piroplasmosis and anaplas- hosts of this species. It is widely distributed in mosis, and Sharif (19.3S) considered this species the Far East and is known to occur in the fol- as a possible disease vector" (Anastos, 1950). lowing areas: Indi;i, Bunna, Ceylon, Indonesia, In Japan, however, its disease transmission capa- Borneo, Malaya, Indochina, Formosa, the Philip- bility is not known. Japanese workers have re- pines, and Japan. Hosts recorded by Sugimoto ported that it attacks humans. 1937a), Nakamura and Yajima (1937), and also ( Genus Boophilus Curtice, 1891 by Kohls ( 1957b ) are usually larger animals such as water buffalo, horse, cow, wild boar, Inornate. Palpi short, compressed, ridged wild pig. goat, dog, tiger, rhinoceros, deer, tapir, dorsallv and laterally. Basis capituli hexagonal and human. dorsally. Eyes present. Festoons absent. The males with adanal and accessory shields. Anal groove obsolete in the female, faintly indicated Amblvomma testudinanum in the male. Caudal process present or absent in the male. Spiracular plate round or oval in ^ Collected and examined both sexes. Coxa I bifid. O Collectpon records (fom literature Type Species: Boophihis annulatus (Say, 1821). ^ Coinljination of above Boophilus microplus (Canestrini) (Fig. 20-23) HaemapIiysaUs micropla Canestrini, 1887:104- 105, 110, PI. 9, Fig. 3, 3a-d, 5, 5a-b. Rhipiceplwhis atinulatus cauclatus Neumann, 1897:413, Fig. 42. Uwboophilm austraUs Fuller, 1899:389-392; Ki- shida, 1939a:541-544. Boophihts annulatus cauclatus: Tokishige, 1911; Ogura, 19.36:75, Fig. 1-6 in Pi. IV.

Boopliilus sp.: Nuttall and Warburton, 1915: 433. BoopfiUus cauclatus: Minniiig, 1934:25-27; Ki- shida, 1927:985; Itagaki, Noda, and Yama- guchi, 1944:1-97, Pi. 15, 19, 1959:1-118, PI. 15, 19; Kitaoka and Yajima, 1958a: 135-147, 1958b: 148-162, 1958c: 179-188. Ihohoophilus sinensis Minning, 1934:25-27; Schulze, 19.3.5:2.^; Kishida, 1939a: .541-544; Nakamura and Yajima, 1937:168-175, PI. XIII. Map 7. Known distribution of Amhhjommii tcsttiili- imriiim. Vroboophilus shariji Minning, 19.34:1-48; Kishi- da, 1939a:541-.544. BlOLOCY: Palpohoc^philus minningi Kishida, 1936:140-142. According to Anastos (1950) all stages in Fig. 1-9; Nakamura and Yajima, 1937:168- the life cycle are known, and this tick occurs on 175, PI. XI. Biological Series, Vol. 15, No. 1 Ticks 31

Margawpits aiitiulatus dtistralis: Sugimoto, there are only three valid species of Boophilus in 1936c: 582-583, 1937b:599-601. the world, but B. kohlsi Hoogstraal and Kaiser, Margaropus annulatus caudatus: Sugimoto, 1960 should be included as the fourth species. 1937h:6()l-603. Although these species are similar in general Palpohoophihis hrachijuris Kishida, 1939a:538- appearance, B. microplus may be distinguished 552, PI. I, II; Shigemori, Aso, and Yajima, by the following combination of characteristics:

1953:290-293. the male has a distinct caudal appendage ( this LJroboopJiilus cdtiihiltis: Ki.shida, 1939a: 54 1-544; serves to discriminate niicro])lus, decoloratus, Nakaimira and Yajima, 1937:168-175, PI. XII, and kohlsi from annulatus), and the dentition of 1942b:34-39, Fig. 1-3 in PI. II; Shigemori, the hypostome is 4/4 in both sexes {micoplus .\,so. and Yajima, 1953:290-293. and kohlsi), while 3/3 or rarely 3.5/3.5 in Boophilus microj)his: Keegan and To.shioka, decoloratus: in the female, the inner margin of

1957:9-10, PI. 8, 9; Kitaoka, 1961a:85-95, the palpal basal segment is concave, while this 1961b:96-104, 1961c: 105-112, 1967:18-21; Ki- segment has an inner bristle-bearing protuber- taoka and Yajima, 1961a:41-52; Kitaoka and ance in decoloratus and kohlsi. Morii, 1963:.32-35, 1967b: 126-129. Hoogstraal (19.56) states that microplus males have adanal shields in which the inner Discussion: margin does not reach the body margin, but in Thi.s tick occurs in Korea, and the Japan, Japanese specimens this spurlike projection often Ryukyu Islands is apparently and the only spe- goes beyond the posterior body margin, and the cies of the genus Boophilus found in these areas. tips may be seen from the dorsal side. It is known under a great variety of names in the Japanese literature. This is partially due to Distribution and Hosts: the fact that most Japanese workers have fol- Anastos (1950) and Hoogstraal (1956) give lowed the classification of Minning ( 1934, 1935, detailed infomiation concerning distribution and

1936). In this paper we have accepted the hosts of B. microjilus. It is known to occur in opinions of Anastos (1950) and Hoogstraal Central America, South America, Australia, the

( 1956) in the classification of species of Boophi- Orient, southern Florida, and Africa. In the lus. These workers reject the subdivisions of the Orient it is known from Fonnosa, Indonesia, the genus and additional species designations pro- Philippines, New Guinea, Borneo, Burma, India, posed bv Minning and are of the opinion that Assam, and small islands of Southeast Asia. Oc- only a careful study of the vast Boophihis ma- casionally it has been found on domestic chick- terials available will resolve current inconsisten- ens and in Africa has been collected from a lion. cies. Thev list three species: B. decoloratus Other hosts recorded are cattle, horses, buffalo,

(C. L. Koch, 1844), B. annulatus (Say, 1821), sheep, deer, and dogs. Kishida ( 1927 ) adds the and B. microj>his (Canestrini, 1887) to which pheasant as a host of this species. must be added B. kohlsi Hoogstraal and Kaiser, Biology : 1960. Of these only microphis has been shown Boophilus microplus is a one-host tick; the to occur in Eastern Asia. Sugimoto ( 19.36c, larvae attach to the host and remain thereon 1937b) reported Margaropus annulatus australis until maturity. Molting and mating occur on and M. a. caudatus ex cattle, horse, and dog the host. "Engorged females leave the host from from Kumamoto, Oita, and Kagoshima prefec- 35 to 149 days after having attached as larvae tures, and also from four islands of the Ryukyus, and there may be from two to three generations hut in the light of our collection these may have a year in South Africa" (Lounsbury, 1905, cited been misidentified. Specimens also have been by Hoogstraal, 1956). "Wilson (1946) observed erroneously identified as Boophilus annulatus no seasonal periodicity of adults in Nyasaland. caudatus l)y Tokishige (1911) and Ogura He found larvae with nymphs and adults on (19.36), both of which were cited bv Sugimoto cattle only once. Nymphs and adults were (19.37b). Kishida (1929) also reported B.^/nni/- ." usually found together. . (Hoogstraal. 1956). latus caudatus from Okinawa and Tanegashima Kitaoka and Yajima (19.58c) calculated the (Is.). The holotype of B. (Palpohoophilus) duration of each stage on bulls which were ex- minningi Kishida, 19.37. as well as other speci- perimentally infested with batches of larvae mens under these various names, were examined throughout the year. "Fifty percent of the bv Keegan and Toshioka, and none of them dif- larvae and nymphs molted 6.6 and 14.0-15.6 days fered significantly from microplus. after infestation, respectively. Fifty percent of Diagnosis: engorged female ticks dropped in 22.1-24.2 days Anastos (1950) and Arthur (1960) .stated after infestation. Adult males migrated to search BmcHAM Young University Science Bulletin 32

Jll' . ^/i*.^<.^>'cCa^

Fio. 20. Boophilus microplus, female. Biological Series, Vol. 15, No. 1 TlCK.S 33

Fig. 21. Bnophiluf microplus, male. Univebsitv Science Bulletin 34 Brigham Young

1 mm

'hi. "TtuaoJ^

Fio. 22. Boophilus micropltts, n\inph. Biological Series, \'ol. 15, No. 1 Ticks 35

0,5 m m

•%.;?t*»«^

Fic. 23. Boophilus microplus, larva. )

36 Bhicham Young University Science Bulletin for females after ecdysis and feeding for 2-3 1958b, 1958c) and Kitaoka (1961a, 1961b, days. The duration of the whole parasitic stage 1961c). was influenced by seasonal temperature to a Rabbits were used as host animals for a nuicli less degree, and the longest duration rearing experiment at the 4()6th Medical Labora- varied from 27-32 days in all seasons in Tokyo." tory. An engorged female ( collected from a A series of excellent physiological studies on this black calf from Ishigaki [Is.], 8 February 1967, species (as B. caudatus and /nicro/j/i/.v) has been Tipton, 67-R-0011)Maid 2,318 eggs from 14 worketl out bv Kitaoka and Yajima ( 1958a, February to 3 March 1967.

Laboratory life cycle of Boophilis microphts ( reared on ears of rabbits

Piuise Date Remarks

Female oviposited on 14 Feb.-3 Mar. Larvae hatched on 17 Mar.-31 Mar. Rate of hatch 98.4% Larvae placed on host 4 Apr. Earliest nymph emerged on 13 Apr. Earliest adult emerged on 21 Apr. Engorged adults dropped on 1 May-5 May 93 females, 78 males through 5 days

Borrelia spp., Coxiella burnetii, Haenuitoxenus velifertis, and NuttaUa equi. Babesia bigemina

is the causal agent of the disease known under the various names of Texas cattle fever, red water fever, splenic fever, bloody murrain, Mexi- can fever, and tick fever.

Genus Dermacentor C. L. Koch, 1844 (Fig. 24-2.S)

Usually ornate. Palpi short, broad, or moderate. Basis capituli rectangular dorsally. Eyes and festoons present. Males with coxa IV much larger than others. Ventral plates or shields ab-

sent in males. Coxa I bifid in both sexes. Spi- racular plate ovate or comma-shaped.

Type Species: Dcnmuvntor rcticuhitus (Fabri- cius, 1794).

We did not attempt to determine the species of the genus Dermacentor for the same reason

that was given by Keegan and Toshioka ( 1957). Whereas adequate infonnation on other genera has been published bv Japanese workers since

1957, verv little is available tor the genus Der- macentor. Kohls (1967) stated that specific detenninations within the genus will continue to be difficult until more extensive studies on in- traspecific variation in the Far East have been made. We followed his suggestion and included Mai* 8. Known distrihntioii of hoophiliis miiroi>lus. only the collection records derived from the literature in chart fomi in Appendix 2. DisE.\sE Relationship: At least four species have been reported as

This species is one of the most important occurring from Korea and Japan bv Kishida pests of cattle, and a vector of organisms patho- (1922a, 19.36), Nakamura and Yajima (19,37), genic to domestic animals in the Americas, Asia, Yajima (1942), and Itagaki, Noda, and Yaina-

and Australia. These pathogens are Aimplasma guchi ( 1944, 1959). They include D. auratus. D. marginiile, Babesia hi<^cmina, B. berbera, B. ovis. coreus, D. recticulatus. and D. variegatus. Arthur BiHLOcicAi. Skiues, Vol. 15, No. 1 Ticks 37

K Tf'^J'^'iAAAAa^

Fin. 24. Dermaccntor sp., female. Brigham Young University Science Bulletin 38

X'^^n^-^j-aA^

Fig. 25. Dcrmtirentnr sp., male. Biological Series, Vol. 15, No. 1 Ticks 39

9>Uy^^»AsJtA.

Fig. 26. Dermacentor sp., female. Brigham Young University Science Bulletin 40

X-^yi-^^yOtji^uy.

Fic. 27. Dennacentor sp.. male. Biological Series, \'ol. 15, No. 1 Ticks 41

Fig. 28. Dennacentor sp., male. 42 BmciiAM VouNG University Science Bulletin

(I960) synonymized D. varieguttis under D. al- Sexual diniorpliisiii sliglit. Palpi usually conical hipictus. with 2nd article salient laterally beyond width The illustrations used in Keet^an and Toshi- of basis capituli (except H. kitaokai). Basis oka (1957) arc reproduced without change, capituli rectangular. Ventral plates absent in the They were made from specimens identified and male. Platelike dorsal spur present on trochan- loaned by Dr. Asanuma (D. auratus and D. ter 1. Spiracular plates usually comma-shaped coreus) and Dr. Yajima (D. reticulatus). in males, round or oval in females. Posterior margin of coxa I never bifid.

Genus Haemavhiisails C. L. Koch, 1844 _ „ ,, i 7. r^ 1 ' •' Type Species: Haenmplnjsam conctnna C. L. Inornate. Eyes absent. Festoons present. Koch, 1844.

Key to the Males of the Genus HaemapJtijsaJis

1. Palpal segment III with dorsal retrograde spur 2 Palpal segment III without dorsal retrograde spur 5

2. Palpal segment III with dorsal spur at middle (Fig. 55 and 66) 3 Palpal segment III with dorsal .spur not at middle (Fig. 38 and 77) 4

3. Palpal segment II with lobelike, anterointemal projection dorsally; ventroposterior mar-

gin fomiing an angle at middle; lateral groove absent (Fig. 55) . hijstncis Palpal segment II without such projection as above; ventroposterior margin slightly convex; lateral groove present (Fig. 66) longicornis

4. Coxa IV with two long .spurs; palpal segment III with a dorsal spur extemolaterally

(Fig. 38) - - -- cornigera group Coxa IV with a short spur; palpal segment III with a dorsal spur on internal margin

(Fig. 77) - wellingtoni

5. Palpal segment II long, clavate, not salient laterally; suture between II and III not clear; spiracular plate about twice as long as wide (Fig. 62) kitaokai Palpal segment II slightly salient laterally with round contour posterolaterally, scarcely beyond basis capituli; suture between II and III well defined (Fig. 47) formosensis Palpal segment II dLstinctly salient laterally beyond basis capituli; suture between II and 111 well defined 6

6. Palpal segment III clawlike, strongly curved inward at tip, overlapping when closed (Fig. 34) '^concinna Palpal segment III otherwise 7

7. Coxa IV with long internal spur 8 Coxa IV with short internal spur 9

8. Color light yellow; length 2.02-2.97 (avg. 2.49 mm), breadth 1.37-1.63 (avg. 1.51 mm);

spur on coxa IV long, sharply pointed, usually straight, approximately Y3 the length of coxa (Fig. 43) jlava Color yellowi.sh-l)rown; length 2.89-3.70 (avg. 3.30 mm), breadth 1.43-2.14 (avg. 1.80 mm); spur on coxa IV stout, wide at base, usually curved, less than half the length of coxa (Fig. 70) vwgaspinosa

9. Coxa I without spur; on Ryukyu black rabbit (Fig. 74) pcntaJagi Coxa I with internal spur 10

10. Posterior half of palpal segment II prominently flared, fomitng bell-shape; junction of segments II and III definitely notched, tarsi with strongly dorsal hump; on dog (Fig. .'30) caiiipanulata Posterior half of palpal segment II otherwise; junction of segments II and III without definite notch; tarsi tapering (Fig. 59) japonica ^(11. iuitmna is not Imiuded in llic almve key.)

Key to the Females of the Genus }IaemaphysaJi,s

1. Palpal .segment III with dorsal retrograde spur 2 Palpal segment III without dorsal retrograde spur 4

2. Palpal segment II strongly salient extemolaterally; palpal segment III with long \entral Biological Series, Vol. 15, No. 1 Ticks 43

spur; hvpostome 4 4 (Fig. 37) , .. cornigera group Palpal segment II moderately salient extemolaterally; palpal segment III with moderate ventral spur; hypostome 5/5 (Fig. 54, 65) 3 3. Palpal segment II with an anterointemal lobelike projection dorsally (Fig. 54) htjstricis

Palpal segment II without such a projection ( Fig. 65 ) longicornis 4. Palpal segment II long, parallel-sided, not salient laterally; comua absent; lateral grooves absent; hypostome 3/3 (Fig. 61) kitaokai Palpal segment not parallel-sided, at least slightly salient laterally; coniua present; lateral grooves present, hypostome 4/4 or more 5 5. Palpal segment II slightly salient laterally with round contour posteroextemally, sairce- ly beyond basis capituli; comua present; lateral grooves present; hypostome 4/4 in anterior half, 5/5 in posterior half (Fig. 46) formosensis Palpal segment II salient laterally, distincdy beyond basis capituli; comua present;

lateral grooves present; hypostome more than 4/4 _ 6 6. Posterior half of palpal segment II strongly salient or flared extemolaterally forming

boot-shape (Fig. 41) or bell-shaped (Fig. 29) _.__ _ .. 7 Palpal segment II salient laterally but not fomiing such shape as above 8

7. Palpal segment II protruding extemolaterally at base at right angle to the axis, boot-

shaped; comua very slight; tarsi tapering (Fig. 41) _ doenitzi Posterior half of palpal segment II prominently flared, bell-shaped; external profile with a definite notch at junction of palpal segments II and III; comua short but well defined; tarsi humped (Fig. 29) campanulata

8. Coxa I without spur; external profile of palpal segment II with a short sahence, pointed, slightly directed extemolaterally at the posteroexternal juncture; on Ryukyu black

rabbit ( Fig. 73) . pentalagi Coxa I with a short spur; external profile of palpal segment II nearly straight or slightly bulged externally at the posteroexternal juncture 9 9. Coxae I-III with short, blunt, subequal spurs; coxa IV with a spur longer than others 10 Coxa I with a spur distincdy longer than others 11

LO. Color light yellow; coxa IV with a spur slightly longer than others; length 1.89-1.92

(avg. 1.90 mm), breadth 1.19-1.28 (avg. 1.2.3 mm)"(Fig. 42) _. flava Color brownish; coxa IV with a strong, triangular spur distinctly longer than others; length 2.99-3.48 (avg. 3.31 mm), breadth 1.88-2.21 (avg. 2.03 mm) (Fig. 69) inegaspinosa

11. External profile of palpal segment III longer than that of II; spurs on coxae II, III, IV short but distinctly extend beyond posterior coxal margins (Fig. 58) japonicn Extemal profile of palpal segment III subequal to that of II; spurs on coxae II, III

blunt, scarcely extend beyond posterior coxal margins (Fig. 33) . ^ concinna

Haemaphysalis campanulata Warburton based on numerous specimens taken from Chi- (Fig. 29-32) nese dogs in Mongolia by M. L. Hearn. He

'''lô ^^'^êd that the type specimens agreed in Haeinapht/salis campamdaia Warburton 1908: "^^^^"^^ ^^* *<^ specimens in the British Mu- 508-514, Fig. 5, 6; Nuttall and Warburton, ^^"'" named H. flava by Neumann. Warburton 1915-491-492 Fig 431-4.32- Kishida 19'?2a- ^908) did not mention the collection data of 852; Nakamura and Yajima,' 19.37:155 PI. VI, ( ^^^ ''^"«'''- ^^^ ^^^ '^^ presumably rs tube No. 1942b:37 Fig. 8 in PI II Fig. 1-5 in PI III- 137 (see Nuttall and Warburton, 1915, 408), A.sanuma, 1942:17-26, 1947b :972, Fig. 2768, p. ^^îch he detemiined to have contamed flava. 1956a: 111-112, 1956b:400, Fig. 225; Itagaki, campanulata, and japonica. Subsequendy, Nut- Noda, and Yamaguchi, 1944:1-97, PI. 4, 1959: Warburton (1915) stated that War- 1-118, PI. 4; Shigemori, Aso, and Yajima, J'^" '''f^ burtons types of japonica (from a tube labeled 1953:290-293; Keegan and Toshioka, 1957:12, "No. 173 H. flava )' were found on Nemor- p] ]7 ^8 haedus crispus, now known as the Japanese Discussion: serow, Capricorni.s crispus, at Hondo, Japan, by Warburton's original description (1908) was the Duke of Bedford's collectors (see discussion

'In Nullall and Warburton, 1915, p. +08, //. Ilava. this number is 1)7. BlUOHAM ^OUNG UnIVEHSITY SCIENCE BULLETIN 44

F. T5UT5U.MI

ft-niiilc. Fig. 29. Ilucmaphijsiilui carnpdnuUita , BlOLOOICAI. ShHIEN. \'()1,. 15, No, 1 Ticks 45

FTSUTSUhl

Fig. 30. Haemaphysalis campamihita, male. 46 HiticHAM Young UNrvF.RSiTY Science Bulletin

F"ic.. 31. Haemaphysulis cainpumilatn. nymph. 47 Biological Series, Vol. 15, No. 1 Ticks

Fic. 32. Haemaphysalis campamdata, larva. 48 BniGHAM Young University Science Bulletin i

of //. japonica). If it is true that all of the specimens in tube N. 173 (No. 137?) were collected from the same host and locality, this collection data of japonica might apply to that of campanulata. Tlicrefore, Warburton's reference to the specimens of campanuhtu, whicli had been mixed with jlata in the tube, might well be the first record of this species from Japan. Nuttall and Warburton (1915) examined Warburton's Mongolian specimens and reported additional samples from Japan. They were collected from dogs in Tokyo and Yokohama and from a house rat in Tokyo.

Diagnosis:

Thi.s common dog tick is distinct from other

haeinaphysalids in that palpal article II is strong-

ly salient laterally; the posterior margin is flared upward to become "bell-shaped," hence the name "campanulata-" the junction of article II and III is deeply notched; the tarsi are humped. Disthibution' and Hosts: This species is known from China, Mongolia. Manchuria, Japan, and Korea. Hosts include cows, horses, dogs, rats, and man. One lot submitted to us (collection, 41-J-(X)2; see Appendix 2) contained .33 females, 11 males, and 4 nymphs, but it seems doubtful that so great a number of ticks were taken from a human host.

There are no records of this species from the Map 9. Known listribution of llucinaplu/sc:li.\ Ryukyu Islands. jxniiiltita.

Biology: stages in the life cycle may be found on the Both immature and adult ticks are very com- ground around dog kennels. Asanuma (1942b) mon on Japanese dogs, even in urban areas. All has provided an excellent account of the life

Laboratory life cycle of Hae maj)]u/ sails campanulata (reared on rabbits) Generation and Stage Ph;i Period in Days Remarks

F, Biological Series, Vol. 15, No. 1 Ticks 49 cvcle of this species. from a horse in Hamgyong-pukto, Korea. Ya-

Rearing was attempted at the 406th Medical jima ( 1942 ) reported it from horses in Aomori Lahoraton'. A single female was obtained from Prefecture. During the preparation of this paper, a dog in Chiha Prefecture, 12 June 1967, and one female and three males loaned by Dr. Kita- 12 females of the F, progeny were fed on oka were available for study. rahbit ears. Pomerantzev ( 1950 ) , who gave a detailed discussion of the morphology, biology, and dis- Disii.^sK Relationship: tribution of the species, states that "it possibly Unknown. covers all of eastern and southeastern Asia (Japan, Kokhinkina)." Haema))In/salis concinna C. L. Koch (Fig. 3:3-36) Diagnosis:

Ihiemaphiisalis concinna C. L. Koch, 1844:237- The male of this species is very distinct in 238; Nuttall and Warburton, 1915: 452-458, that the tips of palpal articles III are curved Fig. 387-393; Kishida, 1922a:852, 1936:142; inward to form "pincers" which overlap when Ogura and Takada, 1927:202-204, PI. XII; closed. Tarsus W is tapered and without a Yajima. 1942:499-510, Fig. 4; Itagaki, Noda, dorsal protuberance. In the female the palpal and Yamaguchi, 1944:1-97, PI. 7, 1959:1-118, profile is very similar to flava and japonica, but PI. 7; Keegan and Toshioka, 1957:13; Asa- may be distinguished from jlava in that the numa, 1965a:lll-112. spur on coxa I is longer than the others, and lliwmaphijsalis Jiirudo L. Koch, 1877:786; Neu- from japonica in that the external profile of mann, 1897:341; Keegan and Toshioka, 1957: palpal article III is subequal to II, palpal article

,30. II has 3-4 infraintemal setae, and spurs on coxae II, III, and IV are wide, blunt and extend lUicmaphijsaUs concinna var. kochi Neumann, scarcely beyond the posterior margin of the 1905:239. coxae. llaemaphijsalis concinna kochi: Newmann, 1911: 111. Distribution and Hosts: The distribution of this species includes Discussion: Western Europe, the Balkan peninsula, and the first referred his speci- Neumann (1897) at USSR. According to Pomerantzev, adults of this mens from Japan to //. liirudo Koch, then subse- species parasitize large domestic and wild ani- quently (1905) renamed them H. concinna var. mals, while nymphs and larvae are found on a kochi, realizing that the type had been lost. The wide variety of small and large mammals and Hil- specimens were three females collected by birds, and occasionally on reptiles. Olsufjev and gendorf Berlin ) and two females from ( Museum Petrov (1960) state that larvae and nymphs are a dog at Saga collected by Yamaguchi. Neu- parasitic on rodents and small mammals. Both variety subspecies, mann (1911) raised this to adults and nymphs have been known to attack but later Nuttall and Warburton (1915), having man. examined Neumann's specimens in the Berlin Museum, svnonvmized the subspecies under Biology: concinna. Koch's original description was not The biology of this three-host tick has been available, but judging from the French quota- studied by Russian workers, and Pomerantzev tion by Neumann (1911)," it is very adequate ( 1950) has briefly summarized the life cycle for detennining the species. As cited by Keegan under laboratory conditions. The engorged fe- and Toshioka' (1957), Nuttall and Warburton male begins oviposition in 4-16 days and lays (1915) condemned the name Jiirudo. Conse- more than 1,000 eggs over a .30-32 day period. quentlv. Neumann's hirudo (1897) might be Larvae begin to appear in 16-30 days and feed the first reliable record of concinna from Japan, for 3-6 days. Larvae molt to nymphs in 16-25 provided we follow Nuttall and Warburton days, and nymphs feed 8-16 days or more. The (1915). cycle can be completed in the laboratory within Ogura and Takada (1927) reported H. con- 120-160 days. Larvae and nymphs can be cinna on cattle and horses at Tokachi, Kushiro, starved for more than 13 months. Pomerantzev Nemuro, and Kitami, ;dl on Hokkaido. Kishida suggests that the life cycle in nature may take (1936) reported collection of this species by M. 1.5-2 years. Mel'nikova (1958) observed the Akasawa in 1928 from North Korea. Itagaki life cycle in nature and concluded that it would et al. (1944, 1959) also recorded a collection take 3-4 years for completion. He also found

= Iic ussim tlors,Tl prosque tide, palpes tourts, jM-'ine plus longc que larf^c — I.. 9 mm; la 7 iimi male inconnu. Brigham Young Univebsity Science Bulletin 50

Fig. 33. Uacmapht/salis coiwinna, female. Biological Series, Vol. 15, No. 1 Ticks 51

/Uotio

Fig. 34. Haemaphysalis conciniia, male. Bhigham Young University Science Bulletin 52

^TL, 7n.i*i>y^

Fig. 35. Ilacnuiphi/salis conciutui, nymph. Bioi.ocicAL Series, Vol. 15, No. 1 Ticks 53

0,5 fTim

'TK Ttu^cJu. 0.7 mm

Fic. 36. Haemaphysalis concinna, larva. .54 BiiicHAM YouNc: University Science Bulletin

rantzev (19.50) described the dentition of the female hypostome as 5/5 or 6/6, but in the specimen examined by us the dentition was 4/4.

Haeinaphijsalis sp. {H. cornigera group) (Fig. 37-40)

llaemaplnjsalis cornigera Neumann, 1897:350- 352, Fig. 16-17; Keegan and Toshioka, 1957: 13, PI. 19, 20; Asanuma, 1965a: 11 1-112, 1965b:.399, Fig. 223. Haemaphiisalis cornigera var. taiwana Sugimoto, 1936a:"l-ll, PI. I-IV; Sugimoto, 1936d:.336-

346, PI. I.

Haenuiplujsalis ias Kishida, 1936: (? unpublished); Nakamura and Yajima, 1937:160, Fig.

10, 11 in PI. VIII, Fig. 1 in PI. IX; Nakatsuji, 1942:295; Yajima, 1950:197-200, Fig. 1-4.

Discussion:

This species is known in the Japanese litera-

ture as H. ias Kishida, 19.36. Actually, it seems that Kishida not only did not publish a description of ias but did not even publish a reference to this name in 19.36. Apparently he had made a verbal reference to the five male specimens

taken from a cow on Miyake ( Is. ) as a new species. In the following year Nakamura and Yajima (1937) published a description of the

Mai- 10. Known cli.stTibulion of Huevuipluisalia con- male of this tick and referred to it as H. ias ciitrui. Kishida, 1936. Yajima (19.50) later described the female and immature stages of the species. that optiinuni conditions for development occur This material was also taken from a cow on in the middle mountain zone ot oak forest and Miyake (Is.). Yajima also mentioned adult ticks inter-alpine valleys (400-800 meters above sea collected from cattle on Hachijo (Is.). Nakatsuji level) covered with broadleaf forests where hu- (1942) reported occurrence of ias on a dog on midity is high. According to Cemy and Kra- Miyake (Is.), but he did not give details. Kee- tochvilova-Kralova (1963), H. concinna in gan and Toshioka ( 1957 ) examined the holo- C/ekoslovakia occurs in southern Moravia and type male and other material loaned bv Dr. southern Slovakia, chiefly on goats. Active Kishida, Dr. Asanuma, and Dr. Yajima. They adults are found mainly in May and June. He stated that all of the specimens were taken from also observed that the entire life cycle took three cows on Hachijo (Is.) and Miyake (Is.) and years. that they were unquestionablv cornigera. Since there is still some doubt about the status of Disease Relationship: H. ias Kishida, 19.36, the Japanese population in This species is reported to be a proven vector this paper is treated tentatively as a part of the of the causal agent (rickettsia) of tick typhus cornigera group until there is opportunitA' to in the USSR and expcrimentidly transmits the further study the problem. virus of spring-summer encephalitis. Pavlovskv and Diagnosis: Soioviyev ( 196.3) demonstrated the capabil- ity of transmission of the virus by bites of the This is the only haemaphvsalid species in the adults as well as transovarian transmission to its Jap;niese fauna in which the male has an excep- progeny. Ticks collected with a flannel-flag har- tionally large coxa IV with two long spurs. bored the agent of tularemia according to Ol- Palpal article II is prominenth" salient laterallv sufjev and Petrov (1960). in both sexes. In the male the ventroposterior margin of palpal article II undulates irregularly, Remarks: giving it the appearance of having two spurs. Nuttall and Warburton (1915) and Pome- The dorsal retrograde spur on palpal article III Biological Series, Vol. 15, No. 1 Ticks 55

F TSUTSUM

Fig. 37. Haemaphysalis sp. (H. comigera group), female. .

Brigham Young University Science Bulletin 36

F.TSUTSUMl

group) Fig. 38. Uaemaphysalis sp. ( H . comigera male. Biological Sebies, \'ol. 15, No. I Ticks 57

0.5 mm \^

'Tn. Tnu.aJu.

Fig. 39. Hucmaphysalis sp. (H. cornigera group), nymph. Science Bulletin 58 R„,n„»v, Y..rNr. Univkbsitv

Fic. 40. Haemaphysalis sp. (//. cornigero group), larva. Biological Series, Vol. 15, No. 1 TicK.s 59

i.s slightly lateral rather than medial. Tarsi III Uaeina))lujsaU.y doenitzi Warburton and Nuttall, and IV have small midventral spurs and api- 1909:64-65; Keegan and Toshioka, 1957:18, PI. coventral spurs. 28; Saito, 1959b: 193-209.

DlSTHlBUTIOX .\ND HoSTS: Discussion: Anastos (1950) discussed the distribution This species was originally described from

and hosts of H. cornigera. It is widely distribut- material collected from a water-hen in Singa- ed in Southeast Asia from Bunna through the pore in 1909. A female tick taken from a pheas- East Indies and Taiwan. It has not yet been ant at Akune, Kagoshima Prefecture, 10 January reported from the Ryukyu Islands, and the only 1956, by Toshioka was the specimen illustrated records from Japan are those from the Izu Is- in Keegan and Toshioka (1957, PI. 28). This lands off the coast of Honshu now included in specimen was listed as species incertae sedis in Tokvo-to. Recorded hosts for cornwera are cat- that manual because Kohls had expressed the tie, deer, water buffalo, pigs, man, and a wide opinion that it was very similar to H. doenitzi variety of smaller mammals and birds. Speci- except for the 5/5 dentition of the hypostome mens reported from Japan have been taken from (original description of doenitzi, 4/4). Hoog- cattle, and on one occasion from a dog. straal ( 1969a ) has expressed the opinion that laboratory rearing will be necessary in order to resolve the problem involving forms in the Ori- ental tropics with 4/4 dentition versus temperate forms with 5/5 dentition.

Diagnosis:

H. doenitzi is a very small species. In the female, the boot-shaped palpal article II is sharply salient laterally and at right angles to the axis of the basis capituli; palpal article III is subtriangular from a dorsal view, without

dorsal spur; the cornua is poorly developed; the hypostome has 5/5 dentition (4/4 according to Nuttall and Warburton, 1915); and coxae I-IV have very short, blunt spurs.

Distribution and Hosts: In addition to the original record of H. doenitzi from a water-hen in Singapore, this species was recorded from Indochina (Touman- off, 1944) without the specific locality or host

being mentioned. In Japan, Saito ( 1959b ) collected immature forms and adults from pheasants and a male from vegetation. In his laboratory he fed H. doenitzi on a domestic fowl, Callus gallus domesticus.

Biology: Unknown.

Disease Relationship: Mai' U. Known distribution of Hacmuphysalis .sp. H. cornigera group). Unknown.

Haenwphiisalis flava Neumann Biology: (Fig. 42-45) Unknown. Haemaphysalis flava Neumann, 1897:333-336, DisE.\sK Relationship: Fig. 3 (in part), 1901:260 (in part); Donitz, Unknown. 1905:129-130; Warburton, 1908:510-512, Fig.

1, 2; Blanchard, 1909:151; Nuttall and War- Haenuiphtjsalis doenitzi Warburton and Nuttall burton, 1915:408-410, Fig. 342, 343; Kishida, (Fig. 41) 1922a: 846-854, 1927:985, Fig. 1897, 1936: 142, Bhicham Younc University Science Bulletin 60

K^>U9«4jJLk.

Fic. 41. Ilacmaphysalis doenitzi, female. Biological Series, Vol. 15, No. 1 Ticks 61

Haemapbysalis tcatanabei Yajima, 1942:500-502, Fig. 1; Keegan and Toshioka, 1957:30.

Discussion:

Neumann's original description (1897) was based on material collected from dog, horse, cow, hare, and vegetation in Japan. However,

as pointed out by Warburton ( 1908 ) , this material (type series) contained at least two distinctly different species. They were H. flava and H. campanulata. Moreover, Warburton

( 1908 ) found japonica mixed with flava and campanulata in a tube labeled as No. 137 "H. flava," identified by Neumann himself, which had been deposited in the British Museum. More confusingly, Donitz (1905) pointed out that H. neunumni Donitz, 1905 (now considered to be a synonvm of H. longicornis Neumann, 1901), was found with flava in the lot which Neumann sent him.

Neumann ( 1897 ) originally described the male of flava as having a long spur on coxa IV, and in 1901 he mentioned the presence of a short-spurred form of flava (Warburton, 1908: 513). Subsequently (1905), he considered his original long-spurred fonn to be a variety {flava var. armata) and in 1911 made the short-spurred fonn the type of H. flava flava. This was criti-

cized by Warburton ( 1908 ) . To make the sitiia- Map 12. Known distribution of Haei}uiphi/salis docnitzi.

tion clear, Warburton ( 1908 ) redescribed the

true flava ( = Neumann's long-spurred form) 1947:972, Fig. 2767; Sugimoto, 1935:23-24. based on one female and one male given him by Fig. 5 in PL I, Fig. 2 in PI. II, Fig. 4 in PI. Donitz, which were taken from a dog at Ise and Ill, 1936c:580, 1937a:308; Nakamura and Ya- from cattle or a horse at Hiroshima. jima, 1937:157; Yajima, 1942:.503-509, Fig. 3; In the same paper Warburton established Nakatsuji, 1942:294; Itagaki, Noda, and japonica ( = Neumann's short-spurred forni) and Yamaguchi, 1944:1-97, PI. 5, 1959:1-118, PI. campanulata (see discussion of H. campanulata 5; Shigemori, A.so, and Yajima, 1953: 290-293; and H. japonica). Nuttall and Warburton (1915) Asanuma et al, 1955:127-128; Asanuma, repeated Warburton's original description and 1956a: 91-96, 1965a: 111-112, Fig. 7.7, 1965b: added measurements and collection records of 400, Fig. 224; Asanuma and Sakurai, 1958: Indian specimens. However, the specimens from 28-39; Keegan and Toshioka, 1957:14-15, PI. India are now considered to be a separate spe- 21, 22; Ohara et al., 1959:8-17, Fig. 1-6, PI. cies, H. indoflava. I-XIV; Saito, 1959b: 193-209, 1960a: 303-321, Kishida 1922a ) described H. i. the holotype 1960b: 189-239, Fig. 1-37, 1962a: 127-146, ( of which was a male collected from a wild boar 1962b: 147-159; Honma and Ohara, 1961:1-7; on Mt. Kiyosumi, Honshu, by Aoki, 30 October Saito, Ohara, and Unagami, 1960: 323-329; 1909. Although the holotvpe specimen was not Saito and Ohara, 1961:1-32; Saito et al., 1965: available for study, both the description and the 14.3-159. illustrations agree with the morphological char- Ilaeuuiphysalis flava var. armata Neumann, acters of H. flava. Asanuma ( 1969 ) concurs with 1905:237-238. this opinion. Haemaphtisalis flava armata: Neumann, 1911: Keegan and Toshioka (1957) examined the Kishida. 112; 1922a:S52. holotype of H. orientally- Kishida, 19.34, taken Ilaemaphiisalis i Kishida, 1922a: 852-853, PI. 26; from a dog at Tokyo, and listed it as a synonym Nakamura and Yajima, 1937:148. of flava. Since the original description was not

Uacmtqyhijsalifi orientalis Ki.shida, 19.34. ( noincn published, orientally is apparently a nomen nuiltmi); Keegan and To.shioka, 1957:14. nudum. 62 BiiicHAM Young University Science Bulletin

Yajiina (1942) described H. ivatanabei based Hjemaphysalis flava oil two females taken from a horse at Okiinaka- yama, Ivvate Prefecture, Honshu, 16 October ^ Collected and examined 1939, by Y. Watanabe. One of the holotype Collection records from series, No. (the O 2535 date cm the label was 16 literature VIII 1939), mounted on a slide, was loaned to (J Combination of above us for study through the courtesy of Dr. Kitaoka,

and it is our opinion that //. wcildiiahei is conspecific with //. flava.

Diagnosis: This species resembles H. japonica but mav be distinguished from that species in that in both se.xes the spur on coxa IV is longer than

the others; in the male the spur is straight, v^)OMivagi sharply pointed, and approximately as long as the coxa; in the female it is slightly longer than ^i\iiigdta(^ '^Fukusfimia\fFukusfiii "^°'^^'9; the other short, blunt, subequal spurs on coxae Shiga /Na^no ^Kvo.o\ />no-, I-III. This species is commonly found on dogs, r^ff"":'!' Shimane YamanashiA: ^ Tokyo whereas there are no collection records of ja- . i, ""fJt^Chiba (J^^^anagawa ponica from dogs. Hirosfiim. (jMivake le .Aichi Distribution and Hosts: y *Tokushima Hachl]o According to the available literature, this ) Kumamoto FMiyazaki species is known only from Japan and Korea. In Japan it has been collected on all four principal islands. Nuttall and Warburton (1915) referred to records from dogs, wild boar (S»/.s- cristatus), and sheep in India; however, Dhanda and Bhat (1968) consider the materials from Map 1.3. Known distribntion of llaemaphijsalia flava. India to be H. indoflava. They state that there hares, is no evidence that H. (//.)' flava Neumann adults were most abundant in late autumn occurs in India. Adult H. flava have been taken and winter. Saito observed a similar trend in most frequently from hares and dogs, but they Niigata Prefecture, except for the fact that there have also been found in considerable numbers was a peak in March instead of winter, and on cows, horses, wild boar, deer, and bear adults disappeared in summer. Nymphs were found active (Ursus thihetanus japonicus). Asanuma et al. on hares throughout the year, but (1955) listed hosts for each stage and indicated generally adults were most abundant from au- that bird hosts were parasitized onlv bv imma- tumn to winter. Larvae were found most fre- ture fonns. Immature fomis are also found on (luently in November in Niigata Prefecture and the same hosts as adults and on small rodents. in summer in Chiba Prefecture. Asanuma (1956a) Saito (1959b) and Saito et al. (1965) al.so studied the laboratory life cycle of H. studied occurrence of this tick in Niigata Pre- flava. Eggs of H. flava hatched in 24-25 days fecture including Sado Island, and they found when maintained at a constant tempeature of fhiva from animals other than those on Asanu- .30 C and at 80-90? relative hmnidity. Eggs de- ma's list. We have identified specimens as flava posited early in May hatched in 28 days at room from the following prefectures: Honshu - Ao- temperature, while eggs deposited in late May mori, Iwate, Fukiishima, Niigata, Ibaraki, Tochi- hatched in .32 days. Duration of larval attach- gi. Tokyo, Kanagawa. Chiba, Shizuoka, Nagano. ment to the host varied from 2-13 days. Most Aichi, Shiga, Kyoto, Nara, Mic, Hiroshima, and lar\ae dropped from the host on the third, fourth, Shimanc; Kyushu - Miyazaki, Kagoshima, and or fifth day after attachment. After de- Kumamoto; and Shikoku - Ehimc and Kagawa. tachment from the host, larvae entered a postparasitic period which varied in length accord- Biology: ing to temperature. At a constant temperature The biology of H. flava has been studied bv of 30 C and at 1005? relative humidity, duration

Asanuma ( 195'6a), Asanuma and Sakurai ( 1958) of the postparasitic period was from 9-15 days in Chiba Prefecture, and by Saito (1959b) in (average 11 days). At 25 C, it was from 14-21 Niigata Prefecture. Asanuma found that on days. When kept at room temperature, after 63 Biological Series, Vol. 15, No. 1 Ticks

Fig. 42. Haemaphysalts flava, female. 64 HiiiGHAM VouNC University Science Bulletin

K.9><^^aA(xhA.

Fig. 43. Hacmaphi/salis fltiva, male. Biological Series, Vol. 15, No. I Ticks 65

0.2 r

Tyi. Triu-o^ 0.2 mm

Fic. 44. Hacnuipht/scili.s jlava, nymph. 66 Brigham Younc University Science Bulletin

Fig. 45. Haemuphi/salK fhivii, larva. ) :

Biological Series, Vol. 15, No. 1 TiCK.S 67 dropping off the ho.st, it was three month.s for the following spring, but as the weather became those detaching in February; for those detach- wanner this period decreased in length. In ing in June, it was 17 days; for those detaching March it was about one month, in May it was in September, it was 17 davs; for those detach- about 10 davs, and in July it was about 6 days. ing in October, it was about 32 davs; for those In general, heavier female ticks deposited more detaching in late November it was 5 months, and eggs than lighter females. Partially fed females the larvae molted the following May. Nymphs were also observed to lay eggs. At a constant remained attached to the host for 2-13 days. A temperature of 30 C the ovipositing period lasted large number fell from the host on the fifth or 12-22 days. The number of eggs produced daily sixth dav. Again the postparasitic period varied reached a maximum several days after the be- in length at different temperatures. It was 16 ginning of the period. From 80-90% of the eggs days under laboratorv conditions, at a constant were deposited in the first half of the oviposition temperature of 30 C and 100% relative humidity. period. At room temperature this period was consider- Asanuma also stated that some ticks which ably longer. For nymphs which engorged in were removed from hosts by scratching or forced February, it was about three months; in July it to leave because of the death of the host before was 16 davs; and in September it was 21 days. they were fully engorged could oviposit or feed

The length of time necessarv for the female to again on a new host. This observation is of engorge varied from 9 to 28 days in several epidemiological importance in that such ticks experiments. Engorged females varied consider- are capable of mechanical transmission of tula- ably in weight. Duration of the preoviposition remia to new hosts. period was 5-90 davs and was correlated with In the 406th laboratory, ticks were fed on changes in temperature. Females which fell rabbits' ears and the following results were from the host in December did not oviposit until obtained

Laboratory life cycle of HaemaphysaUs flava (reared on rabbits)

Generation and Stage Phase Period in Days Remarks

P Adult Feeding 7-11

P Adult Postparasitic

( Preoviposition P Adult Oviposition

F. Egg Incubation

F, Larva Feeding

F, Larva Postparasitic (Premoldng) F, Nymph Feeding F, Nvmph Postparasitic

( Premolting) 68 Bbigham Young I'niversity Science Bulletin

Asanuina ct al. (1955) carried out a series of female, probably from either Ili/strix bengalensis experiments to determine the effectiveness of or Urstis torquattis, in Bunna. Neumann's (1913) //. flava as a vector of tularemia. Adult and record was from dogs in Taiwan. Toumanoff immature //. flava were fed on rabbits infected (1944) reported //. formosensis collected from with tularemia. Ticks which had fed were deer in Haut-Chlong, Indochina. pooled according to stage of development and sex, and material from each pool was inoculated into mice and guinea pigs. AH experimental animals became infected with tularemia, and the etiological agent was recovered from the heart blood of each of these animals. Positive results were also obtained with adults which were infected as nymphs and with nymphs which had been infected as larvae. Studies on transovarial transmission were not attempted. Saito (1959a, 1962b) removed ticks from wild hares which had died of tularemia and found them to be infected with Pasteurellu ttiki- rensLs: He also observed that infected ticks were able to transmit P. tularensis from host to host, but he could not isolate the causal agent of tularemia from ticks collected in the field by the flagging method.

HaemaphtjsaJis jormosensis Neumann (Fig. 46-49) HaemaphijsaUs formoscmis Neumann, 1913:135- 1.37; Kishida, 1922a: 852.

Discussion: This species has not been previously reported from Japan, Korea, or the Ryukyu Islands, but

Neumann (1913) and Ogura (1936) reported it from dogs in Taiwan, and Sugimoto (1936c) cited these records. Other papers by Sugimoto published in 1935-19.37 include records from A/ah 14. Known distrilnition of Hiicnuiplii/niilit jonno- Taiwan but not from Japan or the Ryukvu Is- scn.sis. lands. Kishida (1922a) included H. jormosensis in his key, but he did not give data on hosts Biology: and locality. Thus, the occurrence of this tick Unknown. in Japan was in doubt until it was collected by 406th personnel from wild boar and the Ryukyu Disease Relationship: black rabbit on Amami Oshima. Unknown.

Diagnosis: Haevuiphijsalis fujisana Kitaoka (Fig. 50-53) This species is easilv distinguished from other haemaphysalid ticks bv the following com- llacmuplujsaUs fiijisano Kitaoka, 1970:73-81, Fig.

bination of characters: the palps are blunt; pal- 1-2.5. pal article II is very slightly salient laterallv but

Discussion : extends scarcely beyond the basis capituli, and the lateral margins are almost parallel, with a The relationship between //. jujisami and round posteroexternal margin; palpal article III other HaciiuiphysnUs species which parasitize

has ;i blunt tip (dorsal view) and a ventral spur cattle in Japan re(|uires additional study.

but no dorsal spur; the hvpostome dentition is //. fitjisdiui is related to //. ronciniui Koch, 6/6 in the male ;md 4 4 in the female. 1844, but the male of //. concinna is e;isily separated from //. fujisana in that palpal segment III

DiSTHIBUTION AND IIoSTS: is chelate. Tlie female closely resembles //. con- Nuttall and VVarburton 1915) collected a cinna in general appearance, but "the palpi ot the Biological Sehies, Vol. 15, No. 1 Ticks 69

1/1/ \l y \jg.

P'lo. 46. HacmaphysalLt fnrmosensis, female. BniCHAM VouNC University Science Bulletin 70

Fig. 47. Haemaphysalui formosensis, male. Biological Sehies, \'ol. 15, No. 1 Ticks 71

0. 2 mm

Fic. 48. Haenuiphysalis formosensls, nymph. BnicHAM Young University Science Bulletin 72

Kic. 49. Ilaemaphi/sdlis formosensis, larva. Biological Series, Vol. 15, No. 1 Ticks 73

Fig. .50. Haemaphijsalis fujisana, male. (After Kitaoka. 1970) Bulletin 74 Brigham Voung University Science

Fig. 51. Jhicmaphij.tnlis fuji.santi, female. ( After Kitaoka, 1970) Biological Series, \'ol. 15, No. 1 Ticks 75

Fig. 52. Haemaphysalin fujisaiui, nymph. ( Alter Kitaoka, 1970) Bricham Young University Science Bulletin 76

Fic. 53. Iltwmaphi/sali.s fiijisdiui. lar\a. ( After Kit. ink, i, 1970) Biological Series, Vol. 15, No, 1 Ticks 77 adult of the new .species have a .small, abrupt Holstein calves near the foot of Mt. Fuji, near basolatcral salience that is lacking in //. con- Fujinomiya, Shizuoka, Japan. ciniui. The iininature form has more salient palpi Biology: than the adult" ( Kitaoka, 1970).

This species is also similar to //. japonica in Kitaoka ( 1970 ) reared the progeny of one

tliat the spur on coxa I is longer than those of engorged female. About 250 eggs were deposit- coxae II-IV. But in both sexes the dorsal view ed during a period of nine days. Eggs began to of palpal segment II differs from U. japonica. hatch in 17 days. Larvae fed for 3 to 4 days and In //. fttjisana, palpal segment II has an external molted II to 18 days later at 25 C and in 8 to

profile uliich is longer (male), and subequal to 10 days at 30 C. Nymphs fed for 3 to 4 days in or slightly longer (female) than that of segment August and 4 to 5 days in September. Nymphs III, and the basolateral (posteroexternal) junc- molted in about 15.9 days (males) and 17.5 ture is more sharplv ridged; vcntrallv palpal seg- days (females) at 25 C. ment II bears 3 infraintemal setae. The spira- Disease Relationship: cular plate of the male is longer than wide and without a dorsal process. Unknown. In H. jafwiiica, on the contrary, palpal seg- Haemaphiisalis Injstiicis Supino ment II has an external profile distinctly shorter (Fig. 54-57) than that of segment III, and the basolateral juncture is somewhat blunt with a dull angle; Haemaphi/salis hi/stricis Supino, 1897:251-253;

N'entrally it bears more (5-6) infraintemal setae. Kishida, 1922a: 852; Sugimoto, 19,37a: 31 1-313,

The spiracular plate of the male is wider than 19.37b: 609; Asanuma, l947b:973. Fig. 2771, long and with a dorsal process. 1965a: 111; Keegan and Toshioka, 1957:15, PI. 23, 24; Kawashima, 1963:103; Hoogstraal, DlSTRIBUTIOX AND HoSTS: Trapido, and Kohls, 1965:467-480, Fig. 1-38. This species has been collected only from Haemaplit/salis bisj)inosa: Neumann, 1901:261- 262; (Not Neumann, 1897). Haemapliiisalis nishhiamai: Sugimoto, 1935:29- 33, Pl.'l-III, 19.36c: 58 1 -582, Pi. II, 19.37a:317, 1937d:287-290, PI. Ill; Nakamura and Yajima, 1937:148.

Hacmaphiisalis iuasakii Sugimoto, 1937:610, PI. III.

Discussion: Sugimoto (1937a, 19.37b) reported the occurrence of //. hiisfricis in Yonakuni, one of the Ryukyu Islands, but made no mention of the host. Kishida (1922a) included this species in his key but gave no collection details. Asanuma (1947b) gave a brief account of this species in the Illustrated Encijclopedia of the Fauna of Japan. A collection by 406th personnel includes four lots from wild boar on Ishigaki (Is.) and Iriomote (Is.) in the Ryukyus, but in more recent surveys this species was collected from dogs, pigs, and the Ryukyu black rabbit on Amami Oshima, Kagoshima Prefecture. H. nishit/amai was originally described by Sugimoto, who examined 13 males taken from a wild boar, a dog, and a man on Taiwan. Subse(|uently (19.37a), he listed an additional male from a dog, and in 1937 he redescribed the male and described the female. He stated (1937d) that the male of nishUjamm was very close to lujslricis but distinct from it in that ventral \I \r 1.5. Known distribntioii of lliiciniiphi/suli.-; jiiji.siinii. palpal segment II has "a fairlv strong 78 Bbigham Young University Science Bulletin

Fic. 54. Hacmnphi/sali.s hi/stricis, female. Biological Series, Vol. 15, No. 1 Ticks 79

Fig. .55. Haemaphysalis In/stricis, male. 80 Brigham Vounc UNivEHsiTi- Science Bulletin

Fic. 56. Hactnaphtjsali.'i hi/strici-i, n\Tnph. Biological Series. Vol. 15, No. 1 Ticks 81

2mm

0.2 m m

Fic. 57. Haemaphijsails htfstricis^ larva. .

82 Bbigham Young University Science Bulletin rotrogrado spine" and "lateral grooves modinm, perate belt extending from the eastern Hiina- hunt, including one festoon" and the first leg is lavas of India to the coasts of Vietnam and Fu- of dark color. However, these nioqihological kien Fro\ince of China, and to Taiwan, Ishigaki, characters are not enough to separate this species and Okinawa. Amami Oshima may be now in- from hijstrici.'!, and his description and illustra- cluded in this belt as a result of recent collections fit litistricis well. Hoogstraal et al. (1965) tions. All previously published records from examined two males and two females of ni.slti- China, Ceylon, Macao, Celebes, Java, Borneo, i/anuii from the Sugimoto collection in Taiwan and the Malay Peninsula are either definitely in and considered them conspecific with hi/stricis. error or highh' suspect. Authentic localities for 87° 130° Sugimoto ( 19.37h) described H. iwasukii ht/.^tricis lie between about and east based on a male specimen taken from the bed- longitude and 29° and 16° north latitude. ding site of a wild pig on Ishigaki (Is.). Al- Adults of hystricis have a fairly wide host though the specimen was not available for study, range. the illustration and description indicate that Hoogstraal et al. (1965) listed hosts as hu- iiLasdkii is probablv a svnonym of liijstricis. mans, domestic dogs, buffalo, wild boars (Sits

Recently. Hoogstraal et al'. (1965) studied the scrofci). tigers, hog-badgers { Arctonyx coUaris), identity of liystricis, and they considered //. Teledu badger {Myclantts imtlticei)s), sambar hvasakii Sugimoto, 19.37 to be a synonym of deer (Cervus unicolor), and the Formosan bark- hijstrici.s. ing deer or muntijac ( Muiifiacus rcevesi micru-

Neumann (1901) erroneously described a nis ) species under the name of hispinosa using speci- The often-(|uoted record of a single female mens of a different species. These specimens from the spiny hill tortoise, Geoemyda s])itwsa, were not those from India, the type locality of (Nuttall and Warburton, 1915:425) in "India" hisj)inos(i. but others collected from China and is considered (juestionable since G. spinosa does Japan. The Japanese lot contained two female not occur in India. specimens (Hilgendorf Coll. Berlin Museum). According to Nuttall and Warburton (1915), Neumann misidentified them, and what he

( 1901 ) described as hhpinosa was possibly lit/s- tricis. The following year Neumann (1902) examined Supino's type of hi/stricis. wrongly sy- nonvmi7.ed his])inosa imder Jitistricis, regarding them to be identical. This was criticized by Warburton (1908) and Nuttall and Warburton (1915), but if the specimens were actually hystricis, Neumann's report ( 1901 ) might be the first collection record of In/stricis from Japan.

Dl.\GNOSIS:

H. Injstrici.'i is similar to longicorni.i in having dorsal and ventral spurs on palpal article III, and palpal articl(> II is salient laterallv. How- ever, it mav be distinguished from /ongifor/iiv by the presence of a dorsal knoblike, anterointemal projection on palpal article II, and a median projection on the ventroposterior margin of palpal article II. In the male there are no

lateral grooves, and in the female the comua is short, the basis capituli is at least twice as wide as long with a depression in the middle of the

porous area, and coxa I has a short spur.

Distribution .vno Hosts: This species has previously been considered to be widely distributed in temperate and tropical eastern Asia and on numerous nearby islands. According to Hoogstraal et al. (1965), the dis-

tribution is restricted to a subtropical and tem- Map 1(1 Known distribution ot Hiicnuiphi/scilis tii/xtriiis. ) )

Biological Series, Vol. 15, No. 1 Ticks 83

Biology: allows attachment and feeding; or else nymphs molt to adults on the host rather than on the Ecologically tlii.s tick appears to he a forest- ground. At the 406th Medical Laboratory inhabiting species which occurs at low to inter- nymphs fed on rabbits' ears dropped off the mediate ele\'ations. Newly molted females often host after they had fully engorged. Seventy have been observed crawhiig on birds by Hoog- larvae collected on vegetation on Aniami Oshi- straal et al. (1965), and it is suppo.sed that they ma were reared to adults on rabbits' ears. These seek a host unusuallv early, perhaps several days adults were used for subsequent life cycle before they reach a developmental condition that studies.

Laboratory lite cycle of Haemaphysalis hijstricis (reared on rabbits) Generation and Stage Pliase Period in Days Remarks

P Adult Feeding 8-9 4 unen gorged females (67-J-0308) were placed on host on 29 Nov. 1967. P Adult Postparasitic 7-8

( Preoviposition P Adult Oviposition

F. Egg Incubation F, Larva Feeding

F, Larva Postparasitic

( Premolting F, Nvmph Feeding F, Nvmph Postparasitic

( Premolting) Bbiciiam Young Uni\ersitv Science Bulletin 84

Fig. 58. llacmaphysalis japonicu, female. Biological Series, Vol. 15, No. 1 Ticks 85

Fig. 59. Haemaphysalis japonica, male. Science Bulletin 86 Brigham Young University

0, 1 mm

Fig. 60. Iliicmaplu/sdlis japonica, nymph. ) )

Biological Series, Vol. 15, No. 1 TiCK.S 87

em China, and they .stated that it i.s distinctly .smaller, narrower in front with straighter sides and lateral grooves, and with a longer and sharper ventral spur on palpal article III. Pom- erantzev (1950) referred to dotigkisi as a subspecies and indicated that it inhabits the USSR. The relationship of japonica to japonica dougla.si

is not clear at the moment, and further study will be necessary to elucidate this problem. Hoog- straal (196S) considers japonicii and japonica douglasi to be identical.

Dr.\CNOSis:

This species is close to //. flava and has been mistakenly reported as H. concinna because of the similaritv of the palpal profile. However, unlike flava. the spur on coxa I is longer than the other coxal spurs in both sexes, and especially in the male, coxa IV does not have such a long, sharp spur as in flava. It is also distinguishable from concinna in that the tips of palpal article III are not so strongly recurved inward to become "pincerlike" in the male; the external profile of palpal article III is longer than that of II, while thev are subequal in concinna; palpal artical II has 5-6 infraintemal setae; and the spurs on coxae II and III are short but apparentlv extend beyond the posterior margin of the co.xae. See diagnosis of H. concinna for comparison with that species. M.'ii' 17. Known di.strihution of Haemaplu/salis japonica. Dlstribution and Hosts: often are along with semi-mature Recorded distribution in Japan is Hokkaido, and found describe .\omori. and Nagano prefectures. It is also ones." Somov and Shestakov (1963) known to occur in Korea and China. Pomerant- the habitat of H. japonica dotiglasi as follows: ". . . inhabits foothills and valleys with rich grass zev ( 19.50) gave the distribution of japonica as Southern Primor, USSR, in addition to the growths and shrub thickets. On exposed slopes above, and the distribution of japonica doiiglasi this tick species rarely occurs." (Translation as the USSR and northern China. Hosts record- through the courtesy of the U.S. Naval Medical ed in Japan and Korea are horses, cattle, Japa- Research Unit No. 3, Cairo, UAR. nese serow, deer, dogs, and on one occasion, one Disease Relationship:

nvmph from a wild hare. Pomerantzev ( 19.50 Not known in Japan. In the USSR, japonica gave the hosts for japonica douoja.si as deer, douhedgehog, cattle, dogs, and virus of Russian spring-summer encephalitis humans. Immature stages were found on small (Pomerantzev, 1950), Rickett.'iia sihericus (Si- rodents, badgers, scjuirrels, deer, dogs, Mitstela berian Tick Typhus) has been isolated from H. sibirica. and birds. japonica dotip^la.si by Russian workers (Somov Biology: and Shestakov, 1963). Unstudied in Japaii. Pomerantzev (1950) HaemaphysaJis hitaokai Hoogstraal briefly cited the seasonal occurrence of japonica (Fig. 61-64) doug^Ia.si in the USSR. "From October to March, HacmapJujsalis kitaokai Hoogstraal, 1969b:211- males are exclusively found on animals, often in 221, Fig. 1-35. large numbers; from March through April, they Haemaphijsalis amhifiiia Neumann, 1906:217 arc joined bv females which parasitize mainly (not Neumann, 1901), 1911:109; Kitaoka and during the 1st half of the summer, are found un- Morii, 1967a: 145-152, Fig. 1-9. til October and cease to be found entirely in the winter. Young stages feed from June to August HaenuiphijsaU.'i inerniis: Nuttall and Warburton, Univehsitv Science Bulletin 88 Brigham Young

Fig. 61. Hucviaphysulis kitaokai, female. Biological Series, Vol. 15, No. 1 Ticks 89

Fig. 62. Haemaphysalis kitaokai, male. Bbigham Young Univehsity Science Bulletin 90

Fig. 6.3. Haemaphtjsalis kiUiokai, nymph. Biological Series, Vol. 15, No. 1 Ticks 91

2 mm

J mm

F"iG. 64. Hiwmaphtjsolis kitaokai, larva. 92 Brigham Young University Science Bulletin

1915:367; Kishida, 1922a: 852; Sugiinoto, ferences between specimens from the Japanese 1937a:313-.317, PI. II, Fig. 7, 8 in PI. VIII; population and H. inermis of Eurasia, and they Keegan and Toshioka, 1957:16, PI. 25, 26; suggested that the Japanese material should be

Iwa.saki ct al., 1964:1-10; T.suchic, Iwasaki, a species separate from valid Eurasian inermis. and Okui, 1966:1-15; A.sanuma, 1965a:lll- They tentatively adopted the name ambigua for 112. Japanese material. Haevuipln/sdlis inermis var. aponomoides War- Hoogstraal (1969) subsequently examined burton, 1913:128-130, Fig. 8; Nuttall and the one female specimen ( Berlin Museum No. Warhurton, 1915:367-369.' 173) and the two female andngita identified by

Neumann ( 1906), and after comparing the new- Discussion: ly collected Japanese specimens with numerous collections of the related Eurasian species, he This unic|uc Japanese species has been called concluded that the Japanese population repre- //. tinibi

Sugimoto ( 1937a ) described a Formosan dorsointemal setae and a single ventrointernal species as inermis based on eight females taken seta on palpal segment II of the male, and these from Cervus unicolor stvinhoii on Formosa, giv- setae number two and one respectively in the ing the distribution of the species as occurring female; the male hvpostome is 2.2 times longer also in (see above). Japan than broad compared with three times longer Keegan and Toshioka (1957) identified the than broad in inermi.s; in the nymphs and larvae Japanese population as H. inerinis, and no fur- the tarsi are moderateb- stout in kitaokai but ther attempt was made to clarify the status of iKirrowly elongate in inermis. and the coxa) the Japanese specimen of inermis var. aponomo- spurs are rudimentary in inermis but discrete in ides reported by Nuttall and Warburton (1915). kitaokai (Hoogstraal,' 1969b).

Hoogstraal ( 1962), having examined the type of inermis var. aponomoides and also the tAvo DiSTlUBMTION ."VNT) HoSTS: Fonnosan specimens (RML lot 16603, off "wild This speci(>s has been collected only in Japan deer," Su-ao, Formosa, sent by Dr. Sugimoto), on deer and the Japanese serow in Mie, Shiga, raised the variety to the species H. aponomoides and Kanagawa prefectures, but more commonlv for the population at high altitudes in the Hima- on cattle in the Sanbe Pasture. Shimane Prefec layas and on Foniiosa, He (1962), however, lure. Yajimas single female specimen was from did not see the female specimen ( Berlin Mu- a cow in N;igasaki Prefecture. There are some seum No. 173) of inermi.^ var. aponomoides. records of inermis or inermis var. aponomoides, On the other hand, Kitaoka and Morii but occurrence of these species in Japan is con- (1967a) indicated that there are structural dif- sidered to be erroneous (as mentioned above). Biological Series, Vol. 15, No, 1 Ticks 93

Biology:

Iwasaki et al. (1964) observed the seasonal occurrence of this tick in a pasture at the foot of Mt. Sanbe, Sliimane Prefecture. There are two Haemaphysalis kitaokai peaks of adult prevalence, one is a small peak appearing early in spring to June and another ^ Collected and examined more p r o m i n ent peak from October to No- O Collection records from vember. No larval and nymphal fomis were literature found in the pasture or on cattle. Kitaoka and (J Combination of dbove Morii ( 1967a) suggest this is possibly due to the fact that the immature forms are exceptionally rapid feeders and are attached to the host for a short period of several hours. Tsuchie et al. (1966) reported observations on overwintering of this tick. On 19 November 1964, they placed jars containing 60 engorged ticks and dried, fallen leaves out-of-doors, and found (|uiescent ticks 1 cm deep under the leaves on 18 Febru- ary the following year. The 53 females which had successfully wintered oviposited in early May. Ten specimens were observed and they laid 109-1,020 (average 617) eggs during the period from 5 May until 6 June, when the average outdoor temperature began to rise to 15 C at the Sanbe Pasture. Kitaoka and Morii (1967a) studied the biology of this species at Kodaira, Tokyo, using material originally collected at Sanbe Pasture in November 1962. They suggested that the life cycle was probably completed in one year in

nature. The following table is a summary of data from Kitaoka and Morii (1967a). Nymphs

Map 18. Known di.stribution of Haemaphysalin kitaokiii. and adults were fed on calves' ears.

Laboratory life cycle of Haemaphijsalis kitaokai

Generation Period in Days and Stage Plia; 25 C Outdoor Remarks p 94 Bmr.HAM Young ITniversitv Science Bulletin on the rabbit on 8 November dropped off on 15 nosa in the Nuttall Collection (N167) in the November and laid 823 eggs from 10 to 30 British Museum and a paralectotype of neu- December from which 588 larvae hatched out manni (N1425) in the Neumann Collection in beginning on 7 January. The F, progeny were the National Veterinary School, Toulouse. The fed on a rabbit's ear on 22 January, dropped off former specimen was taken from southern on 23 January, but all died before they molted. Madras, India, and the latter was from Japan. Another lot of F, larvae were put on a rabbit's Since then it has been thought doubtful that ear at 1400 of I Fefiruai-v and were observed to l)isj)inosa actuallv occurs in Japan. .All materials have already dropped off by 0800 the next in the 406th collection have been examined by moniing. The rapid feeding of larvae was also Kohls and Hoogstraal and identified as H. neti- observed, but only a single larva survived until manni. It is likely that the specimens in the it molted to a nvmph on 3 March. The room Japanese literature are also netimanni, since our temperature in the laboratory was approximate- specimens were collected from the same host ly 25 C. species and in essentially the same geographical

localities. More recently Hoogstraal al. Disease Relationship: at (1968) compared numerous specimens from Unknown. Australia, New Zealand, tropical Asia, northern Haemaplnjsalts longicornis Neumann China, Japan, and Korea. They confinned that (Fig. 65-68) specimens of "bispinosa" and "netimanni" from temperate northern China, Japan, Korea, and IIaei)ui))luj.siilis longicornis Neum;mn, J901;261, northeastern USSR are structurallv identical to Fig. 2;' Hoogstraal, Roberts, Kohls, and Tip- those from Australia and New Zealand referred ton, 1968:1197-1213, Fig. 1-36. to as H. longicornis Neumann, 1901, which ap- Haenuiphtjsalis netimanni Donitz, 1905:127-129, pears to be a prior name applicable to these Fig. 4-6; VVarburton, 1908:508-519; Neu- northern populations. We accept their opinion mann, 1911:109; Kishida, 19.36:139, 1939b: and use the name H. longicornis in this paper. 17-19, PI. 4; Trapido, 1965:160; Hoogstraal Published reports indicate there are tvvo strains and Trapido, 1966:1192, Fig. 10-26. (bisexual and parthenogenetic) within the Japa- Ihwnmphiisalis bispinosa (not Neumann, 1897); nese population. H. ])is])inosa Neumann, 189"?, a Nuttall and Warburton, 1915:426-4.33, Fig. tropical Asian species, tvpically a bisexual spe- 358-362 (in part); Kishida, 1922b:8.52; Ogura cies, is known from India, Ceylon, Pakistan, and Takada, 1927:204-205, PI. 14; Sugimoto, Nepal to Bumia, Thailand, and the Malay 1936c:580, PI. I, 1937b: 605-609, PI. II; Naka- Peninsula. mura and Yajima, 1937:151, PI. IV, 1942a:21- 33, 1942b:.34-39, PI. II, III; Nakatsuji, 1942: Diagnosis:

294-295; Yajima, 1942:,502-509, Fig. 2, 1963: H. longicornis is very close to H. bispinosa 103; Itagaki, Noda, and Yamaguc^ii, 1944:1- Neumann, 1897 but may be distinguished from 97, PI. 3, 1959:1-118, PI. 3; Asanuma, 1947a: that species on the basis of its larger size: male 7-14, 1947b:97.3, Fig. 2770, 1958:279, 1965a: 2.47 mm (2..30-2.57); female 2.60 mm (2.0-2.8); 111-112, Fig. 7.6, 1965b:399, Fig. 222; Shige- while in bis))inosa. male, 2,0 mm; female, 2.2 mori, Aso, and Yajima, 1953 :290"- 293; Namba, mm in length; hvpostome with dentition of 5 5 1953:130-1.3,5, 1954:49-55, 1958:1-99, PI. I, (rarely 4/4 in female), bispinosa always with 196.3a:21-28, 1963b: 103-114; Chikaki, Otake, 4/4 dental formula; UMnphs with 3/3 dental and Miiira, 1936: .53-62; Keegan and Toshi- fonnula, whereas bis])inosa with 2/2; nymphs oka, 19.57:11-12, PI. 1.5-16; Kitaoka and Ya- without dorsal spur on palpal article III, but jima, 195Sa: 1.35-147, 195Sb: 149-162, 1958c: bispinosa has a bulging posterodorsal margin of 179-188; Kitaoka, 1961a:S5-95, 1961b:96-104, palpal article III. //. longicornis may also be 1961c: 105-1 12, 1961d: 142-144, 1962:106-111; distinguished from H. hystriris on the basis of Nagahana and Mat.suo, 1962:119-121; Saito the following characters: well-de\el()ped comua; and Ohara, 1961:1-32, Fig. 19, 20, 26; Saito, no dorsal knoblike projection on anterointenial

1962a: 127-146, Fig. 1-30;" Saito ct al., 1965: margin of palpal article II; lateral grooves pres- 143-159, Fig. 1-24. ent in the male and shape of spiraciilar plate as illustrated. Discussion:

For many years this tick has been referred DlSTBIBUTIOX AND HoSTS: to as //. bispinosa Neumann, 1897 in the Japa- Neumann's original description (1897) was nese literature. Hoogstraal and Trapido (1966) based on females from a cow. Kempsev. New selected and studied a female neotype of Jmpi- South Wales, .Australia. .According to Hoog- 95 Biological Series, Vol. 15, No. 1 Ticks

Fio. 65. Hacmaphi/salis Inngicornis, female. 96 BniGHAM Young UNrvERsrrY Science Bulletin

Fit;. 66. Haenmphijsah.s longicornis, male. BioLiKiicAL Series, Vol. 15, No. 1 Ticks 97

Fig. 67. Haemaphtjsalis longicomis, nymph. Bulletin 98 Bhicham Vounc Univehsitv Science

0.2 m m

0.2 mm 9n. '^TUoaAi.

Fig. 68. Hacm(ii>)uistilix /orifjifonii,?, larva. . )

Biological Series, Vol. 15, No. 1 TiCK.S m straal et al. (1968), this species occurs in Aus- geographical locahty, and our collection con- tralia, New Zealand, New Caledonia, Fiji Is- Finns Kitaoka's finding ( 196Id) of predominant- lands, Tonga, Friendly Islands, Efate Island, ly parthenogeiietic populations in the north of New Hebrides, northern China (Peking), Japan, Japan and bisexual populations in the southern Korea, and northeastern USSR. They suggest islands. However, we have both parthenogenetie that the bisexual population is presumably pre- and bisexual populations from Oki Island. In dominant on the Korean peninsula. It may be the laboratory the bisexual strain readily con- noteworthy that reliable records of this spe- verts to parthenogenetie type of reproduction cies on the R)ukyu Islands cannot be found. The when the females are isolated from the males, distribution and sex ratio of this species in Japan Bremner ( 1959) found a ratio of approximately have been reviewed by Kitaoka (I96Id). I male to 400 females in the Australian popula- H. longicornis parasitizes many animals from tion and no spermatozoa in all males dissected, small to large mammals, and birds. Hosts re- irrespective of whether they were fed or unfed, corded are cattle, horses, sheep, goats, deer, Excellent physiological and ecological studies bears, pigs, foxes, racoons, badgers, cats, dogs, of this species (as bispinosa) were done by rabbits, house sparrows, skylarks, thrushes, tiir- Kitaoka (1961d), Namba (1953, 1954, 1958, kevs. ducks, chickens, pheasants, and humans. 1963a, 1963b), and Asanuma (1947a). Naka-

mura and Yajima ( 1942a ) studied the laboratory life cycle. In the 406th Medical Laboratory both iiiOLOG\ parthenogenetie and bisexual strains were reared

This tick is known to be parthenogenetie. In on rabbits' ears, and their life cycle data is sum- Japan the ratio of males to females varies with marized in the following table.

Laboratory life cycle of Haemapluisalis longicornis bisexual populations ( parthenogenetie and (reared on rabbits) Period Number of Days Least Most Average P B P B P B

Oviposition to hatching 24 38 31 39 27.5 38.6 Resting (larvae) 3 3 5 4 4.0 3.5 Feeding (larvae) 4 5 5 7 4.5 6.0 Drop-off to molt (larvae) 14 20 17 22 15.5 21.0 Resting (nymphs) 2 3 3 3 2.5 3.0 Feeding (nymphs) 5 4 7 5 6.0 4.5 Drop-off to molt (nymphs) 12 14 16 16 14.0 15.0

Resting (adults) . . 4 2 6 4 5.0 3.0

Feeding (adults) . 11 11 19 13 15.0 12.0 Drop-off to oviposition (females) 3 3 7 6 5.0 4.5 Oviposition (females) 11 17 27 18 19.0 17.5 Number of eggs 2,024 2,740 Egg hatch (fc) 94 96

P = Parthenogenetie population B — Bisexual population

Disease Rel-^tionshu': Re.marks;

This species is a proven vector of Coxiella The only known example of triploidy in ticks hiirnclii (Q fever) among Australian cattle and occurs in //. lonoicorni.s. Oliver and Bremner

is also capable of transmitting the causal virus ( 1968 ) found the chromosome number to be 32

of Russian spring-summer encephalitis in north- or .3.3 in the obligate parthenogenetie female and

em USSR. In Japan it is suspected of trans- 31 in the reproductively nonfunctional male in mitting bovine piroplasmosis. Experimental in- the Australian population. According to infor- fection of calves with Theileria nmtans by the mation supplied by Oliver and Tanaka (1968), bite of this species (listed as bispinosa) was the chromosome number of the Japanese par- reported by Ishihara and Ishii (19.56). Namba thenogenetic population is 32 or 33 in females (1963b) also referred to this species as an im- and 31 or .32 in males (t^vo males found among portant vector of T. nmtans in Japan. approximately 3,000 samples of females which 100 Bhiciiam Young University Science Bulletin

specimens were loaned for study through the Haemaphysalis longicornis courtesy of Dr. Kitaoka, and two females were found in Dr. Yajima's collection at the National ^ Collecled jnd examnied Institute of Animal Health, Kodaira, Tokyo. O Collection records from literature

(^ Combination of above Diagnosis: in , Abash Kamikawa_ This species is close to //. fhiva in general Ishfkar appearance but may be distinguished in that the Shinbesh) HWr-V^^ Tokachi // body color is brownish and darker, and the body Hiyania •V^'bur_ size is remarkably larger in both sexes. In the Oshima

'( male, the conuia is longer; the dentition of the iAomori hvpostome is 5 5 to 6/6; the posteroexternal

•,'C^lwaTe juncture of palpal segment II is somewhat blunt, whereas in flava the angle is sharper, the juncture itself being a small tip; legs more robust; NiigalaQ/ trochanteral spur I is much reduced; coxae I and / flQpukushimaoQFukushii (^mGumma Tochigi IV have relatively shorter spurs, especially on coxa IV the spur is stronger, less #Tottor, (^"^''O.^S^'"'"'' more or curved, ^-^., ^\*-' ^ TokyoOQChiba wider at base; tarsi II-IV have a triangular, spurlike ventral projection at approximately the mid- Tsushima y'riJ^Q^C^^ C\h,z^^Q 0,h,m3 H.joshimaMto-ykayama W H,,,:, dle. The female of this species is quite similar lki*^£j:ukuoka 7^S V to that of flava. However, palpal segment II ji"<^^0,tay/ Tokushima Hach,|oko#^j ^ C^^0j5'^'"^'"o*o Hachijo differs ventrobasally as in the male (see illustra- NagasakCq'lVI,y^^3l^, tions), and coxa IV has a longer spur. Aogashima^ Kagoshima

•QO Distribution and Hosts:

Saito collected H. megaspinosa from vegetation, but adults of this species were collected

Mat 19. Known clistribiitioii of Haciiuiphi/salis lon'^- from a wild boar, deer, and Japanese serow

icornis. {Capricornis crisjnts). This species is known at pre.'ent from Mie, Shiga, Miyazaki, and Kanaga- wore produced from ten batches of eggs), and wa prefectures. Immature forms have not yet in the Japanese bisexual population it is 22 in been collected from the field. females and 21 in males. In parthenogenetic females from Cheju Do, Korea, 22 to 24 chromo- Biology: somes have been observed. This species fed on rabbits in the 406th Med- ical Laboratory', and the life cycle was studied. Haenuiphysalii megaspinosa Saito Compared with //. it showed slower de- (Fig. 69-72) flava, \elopment in each stage. Five engorged females Uiiemaphiistilis uiegospinosa Saito, 1969:87-96, (6S-J-0301 ) began oviposition on 29 to 31 March Fig. 1-16. 1968 and laid batches of .368-2,077 eggs during the ovipositing period of 23 to 26 days. After the Discussion: incubation period of 33 to 37 days, larvae Saito. using the flagging method, collected hatched out on 3 to 6 May. The hatch rates of adults of this species in a cedar forest of a moun- the two batches were 98.1^ and 96.9?. A con- tainous area in Kanagavva Prefecture. Personnel siderable number of larvae from the two colo- of the 406th Medical Laboratory collected this nies were fed on a rabbit for life-cycle studies. species from a wild boar, deer, and Japanese In the first series of rearing experiments, larvae serow. Three lots. J-()177, J-()179, and J-01S4, were put on the host 9 days after h;itching and collected from a wild boar and two deer in Mie nvmphs were put on the host 4 davs after molt- and Shiga prefectures, contained this species ing. In the second series, larvae were put on along with //. jlava. The female of H. mega- the host 2S days after hatching and the nymphs spinosa resembles //. flava: therefore, the rec- were put on th(" host 26 davs after molting. The ords of fhivii from thes(> hosts which have been feeding periods of larvae and nvmphs did not previously reported by Japanese acarologists differ essentiallv between the two experimental might possibly include H. megaspinosa. Other series, and the results are sumnnirized as follows: Biological Series, Vol. 15, No. 1 Ticks 101

0.5 mm O.I mm

' Am

Fig. 69. Haerrmphysalis megaspinosa, female. 102 Bricham Young University Science Bulletin

2 mm

/W:^H

r. /\^i<,

V\c 70. Uui'muphysalix rneguspinosa, male. Biological Series, Vol. 15, No. 1 Ticks loa

^TTL nti^. 0.2 mm

Fig. 71. Uacinapliysalis megaspinosa, nymph. 104 BiucHAM YouNC University Science Bulletin

Fk;. 72. Ilacmaphiisuli'i iiiei^titipiuosa, larva. BiDi.otiicAL Series, Vol. 15, No. 1 Ticks 105

Hoog.straal and Yamaguti,

M.\p 20. Known distribution of Hucmupht/sidis lucg^ii- spinosa.

Laboratory life cycle of Haeinaphysalis megaspinosa and Haeniaphijsalls flava

Period in Days

Stage Phase megaspinosa flava

Adult Bbigham Younc University Science Bulletin 106

77 A^

Fk;. 73. IhKiniiphij.salix pctitiiltif:,!, female. Biological Series, Vol. 15, No. 1 Ticks 107

r A^4o

Fic. 74. Haemaphijsalis pentalagi, male. Brigham Young University Science Bulletin 108

Fig. 75. lUwmaphysalis pcntahigi, nymph. Biological Series, Vol. 15, No. 1 Ticks 109

-^. 7K,.*

Fig. 76. llaenuiphysulis i)entaUigi, larva. 110 BniGHAM VouNc University Science Bulletin wide spurs which arc hardly discernible; there Distribution and Hosts: is no dorsal spur on palpal article III; palpal Known distribution is Amami Oshima in the article II has a pointed, externally directed pos- Uvukvu Islands. Pospclova-Shtrom's record may teroexternal juncture; in the female, the capitu- be trom the same island. The only known host luiii and scutum are very large in relation to is the Rvukvu black rabbit, Pcntahi

Laboratory life cycle of UdemaphijSdlis pentalagi (reared on laboratory white rabbit) Generation and Stage Phase Period in Days Remarks

P Adult Feeding

p Biological Series, Vol. 15, No. 1 Ticks 111

Fig. 77. Haemuphysalis wetlingtoni, male. 112 BiiK.iiAM VouNG University Science Bulletin

Disease Relationship: groove curved around tlie anus anteriorly. Spi- racular plate oval. Sexual dimorphism IJnknovvn round or marked. Venter of male eoveretl with seven Genus Ixodes Latreille, 1795 plates.

Inornate. Eyes and festoons absent. Anal Type Speciks: Ixodes riciniis (Linnaeus, 1758).

Key to the Males of the Genus Ixodes'

'2 1. Coxae without internal spur Some coxae with internal spur 4 2. Legs distinctly longer than body; hypostome pointed at tip; on bats (Fig. 116) vespertilionis

Legs not distinctly longer than body; hypostome blunt or notched at tip - 3 3. Anal grooves convergent; on sea birds (Fig. 105) ^ signatus Anal grooves divergent or parallel; on swallows (Fig. 87) Uvidus

4. Coxa I with one spur 5

Coxa I with two spurs , 6

5. Coxa I with a long internal spur; on badger (Fig. Ill) tanuki

Coxa I with a short, blunt internal spur (Fig. 96) ovatus

6. Coxa I witli a thick, long external spur acutitarsus

Coxa I with a blunt, short external spur 7 7. Hypostome with moderate denticles; internal and external spurs of coxa I short, subecjual (Fig. 83) '^ranuJatus Hypostome with large, strong denticles laterally; internal spur of coxa I longer than

external spur . 8 8. Spiracular plate ellipsoidal, much longer than wide, usually anglelike contour antero- ventrallv; mostlv from western part of Japan (Fig. 92) nipponensh Spiracular plate oval, longer than wide; mostly from northern part of Japan (Fig. 100) persulcatus

^Ixodes philipi is not included in this key.

Key to the Females of the Genus Ixodes

\. Coxae without internal spur - 2

Some coxae with distinct internal spur - 3

2. Anal groove pointed in front; posterior amis of anal grooves divergent; on bats ( Fig. 108) simplex simplex Anal groove rounded in front; posterior arms of anal grooves convergent or subparallel 3 3. Legs distinetlv longer than body; hypostome lanceolate; scutum widest behind middle; on bats (Fig. 115) vespertilionis Legs not distinetlv longer than body; hypostome blunt at tip; scutum widest at middle or in front of middle; on birds 4 4. Auriculae well developed; on sea birds 13 -Auriculae a1)sent; on swallows (Fig. 86) Uvidus

•5. Coxa I with one spur 6

Coxa I with two spurs 8 6. Coxa H with a short external spur; scutum with posterolateral margin slightly concave (Fig. 90) monos])inosus Coxa H without an external spur; scutum with posterolateral margin convex 7 7. Coxa I with a long internal spur; on badger (Fig. 110) — tantiki Coxa I with a short internal spur (Fig. 95) ovatus

8. Auriculae well developed - 9 Auriculae absent or poorlv developed 11

9. Internal spur on coxa I much longer than external spur; auriculae nonnal 10 Internal spur on coxa I nearly e(]ual to external spur; auriculae well developed laterally (Fig. 112) ' ' turdus

10. Internal spur on coxa I sharply pointed, long, iisuallv ovcrlipping on approximately /a of coxa II; postscutal marginal bodv setae pointed, length approximately 0.08 mm; HioLOCMCAL Series, Vol. 15, No. 1 Ticks 113

inastly from northern part.s of Japan (Fig. 99) persulcatus Internal .spur on co.xa 1 long, u.sually overlapping anterior margin of coxa II (Fig. 80); postscutal marginal body setae scaled, forked, length approximately 0.14 mm; mostly from western parts of Japan (Fig. 91) nipponensis 11. Coxa 1 with a thick, long external spur; scutum cordate, nearly as wide as long (Fig. 78) — acutitarsus Coxa I with a blunt, short external spur; scutum distinctly longer than wide 12 12. Anal groove slightly pointed in front; genital aperture on level with coxa III (Fig.

79) __ angustus Anal groove round in front; genital aperture on level with coxa IV (Fig. 82) granulattts 13. Dorsum except scutum hirsute, uniformly covered with white hairs; basis capituli subtriangular dorsally; porous areas well separated (Fig. 10.3) philipi Dorsum except scutimi not hirsute; basis capituli rectangular dorsally, much wider than long; porous areas very close to each other (Fig. 104) signatwi

gan and Toshioka, 1957:18-19, PI. 29; Ohara and Tamura, 1958:2.3-24. EscJiatocephalus acutitarsus: Neimiann, 1901: 290. Ixodes happinus: Keegan and Toshioka, 1957:31.

Discussion: Nuttall and Warburton (1911) discussed the synonymy of this species and mentioned that Karschs type came from Japan. These workers also stated that they had seen two females from Japan and Formosa. Sugimoto (1936b, 1936c) merely cited records of Nuttall and Warburton (1911). Kishida (1930a) reported occurrence of this species on the serow, Capricornis crispus, in Gifu, Nagano, and Toyama prefectures, Japan,

and later ( 1935 ) from cattle on Formosa. Asa- numa cited the records given above and added the dog and the cow to the host list in the 1947 edition of the Illustrated Encijclopedia of the Fauna of Japan. Keegan and Toshioka (1957) examined two loaned specimens taken from man, one from Wakayama Prefecture and the other from Saitama Prefecture, and identified them as acutitarsus. Yajima's (1955) record was from a cow in Iwate Prefecture, and he reported this tick merely as n. sp. and did not provide a specific name. This specimen was identified by Mr. G, M. Kohls as acutitarsus. Keegan and .M.\p 22. Known distribution of UaciiKijihi/.'idlis wcll- in^toni. Toshioka (1957) referred to this specimen as Ixodes happinus, but this is an error since Dr. Ixodes acutitarsus (Karsch) Yajima had not given a name in his original (Fig. 78) description, nor used this name in any publication since that time. Ohara and Tamura (1958) Haemalastor acutitarsus Karsch, 1880:141-142; reported this tick on man. Neumann, 1899:180.

Ixodes acutitarsus: Neumann, 1901:286; Nuttall DiACNOSIS: and Warburton, 1911:202-203, Fig. 195; This tick is said to be the largest in the that Schulze, 1935:2.34; Kishida, 19.30a:2, 19.3.5: genus Ixodes. Asanuma ( 1947b ) stated 137; Sugimoto, 1936b: 1-20, 1936c:585; Asa- even unengorged females measure as large as numa, 1947b:972, Fig. 2766. 1965a: 113- 116, 8 mm in length. It mav be distinguished from 1965b:399, Fig. 220; Yajima, 1955:52-53; Kee- other Ixodes spp. by the following characteris- 114 BniGHAM Young University Science Bulletin

Fig. 78. Ixodes acutittnstis, female. Biological Series, Vol. 15, No. 1 Ticks 115

tics: coxa I i.s deeply inci.sed witli two thick, long 1952:111-116; Asanuma, 1955:1240-1242. spurs; there are neither conuia or auriculae on Discussion: the basis capituli; the anal and genital grooves extend dorsallv to reach the distinct lateral In the 1947 edition of the Illustrated Ency- groove of the dorsum, thus giving a iestoonlike clopedia of the Fauna of Japan, Asanuma appearance; the legs are slender, the tarsi gradu- (1947b) recorded the occurrence of this tick in ally taper distally, especially in the teinale. Sakhalin and in 1951 gave a description of females taken from the red-backed mouse, Cleth- DiSTniBUTION AND HoSTS: rionomys rufocantts (Sundevall, 1846), in Sak- Nuttall and Warburton (1911) recorded col- halin. Asanuma and Sekikawa (1952) and Asa- lections from Japan, Formosa, and Sikkim from numa ( 1955) reported immature forms of Ixodes Japanese serow, dog, cow, ;ind human. All species on rodents from Hokkaido, and they specimens were adults; innnature forms have temporarily numbered these forms as sp. 3 for never been collected. the larvae and sp. 61 for the nymphs. Later, Asa-

numa ( 1957 ) again reported these forms from Hokkaido and stated that the previously report- Ixodes acutitarsus ed sp. 3 and sp. 61 were the immature stages of Ixodes angiistus. 9 Collected and examined O Collection records from Diagnosis: literature (^ Combination of above The female of this species is close to /. granulatus in the key but may be distinguished from that species in that the anal groove is slightly ogival in front; the genital aperture is on a level

with coxae III; the hypostome is shorter than the palps and sharply pointed at the tip, the denticles of the inner file are subequal to those of the outer file and occur on almost the entire dentate length except two or three rows near the tip, while in grantdatus the denticles of the inner file are much smaller than those of the outer file and do not occur on the proximal two- fifths of the dentate length; the scutum has well- defined, straight carinae, and the lateral margin of the scutum has a distinct angle which sepa- rates the anterolateral and posterolateral margins. According to Asanuma (1951), this species

is also allied to /. soriciis, I. muris, I. jellisoni,

and /. minor but may be distinguished from them by the smaller size and wrinkled integument of the postscutal area, the dentition of the hypostome, the absence of comuae, the larger porous areas, and the strong lateral angle of the scutum. Map 2.3. Known distribution of Ixodes uciititursut;. Distribution and Hosts: Biology: Known distribution of this tick includes North Unknown. America, Canada, Alaska, Sakhalin, and Argen- tina. Hosts recorded are rodents, other small DisE.\SE Relationship: mammals, and sometimes humans. Unknown. Biology: Ixodes angtisfus Neumann Little is known about the biology of this tick. (Fig. 79-81) Males are rarelv found on host animals, and Ixodes anoti.stus Neumann, 1899:136; Asanuma, copulation probably occurs in the nest of the 1947b:97I, Fig. 276.3, 1951:1-4, 1957:2:34, host. 1965a:113-116," 1965b:.398, Fig. 219; Ono, 1966:62-68. Disease Relationship: positive rec- Ixodes sp. 3 and sp. 61 Asanuma and SekikawM, Cooley (1946) reported three 116 Brigham Young University Science Bulletin

/ \

• A.

Rvjuihi

Fig. 79. Ixodes aniiufihis, female. BionK-.icAL Series, Vol. 15, No. 1 TlCK.S 117

7n.. TKic^xM, 0.2 m m

Fig. 80. Ixodes angnsUis, nymph. Brigham Young University Science Bulletin 118

0,2 m m

0.1mm ^. oAt.Xoo-A-

Fic. 81. Ixodes tmnustus, larva. Biological Series, Vol. 15, No. 1 Ticks 119

mens were actually grcinuhitus. Arthur (1957) Ixodes angushis discussed Korean specimens.

^ Collected and examined Diagnosis:

O Collection records from /. gramdattis is easily distinguished from literature other Ixodes species by the following combina- ^ Combination of above tion of characters: coxa I has two short spurs, the internal spur is slightly longer than the external spur; the cornua is very short in the male and absent in the female; the auriculae are poor-

ly developed; the scutum of the female is finely granulated, longer than wide, covering more than half the length of the dorsum and the scapulae are pointed.

Distribution and Hosts: Anastos (1950) cited distribution records from India, Burma, and the East Indies. Kohls

( 1950 ) gave a collection record from Luzon Island in the Philippines and (1957b) from Ma- laya. Collection records indicate that this tick is primarily a parasite of rodents. Asanuma and

Kosaka ( 1955) took a female of this species from the thrush, Turdits celaenopus celaenojnis, on Miyake (Is.), Tokyo-to. Prior to our survey, material in the 406th Medical Laboratory collections was from the Ryukyu Islands.

Ixodes granulatus Map 24. Known dLstribution of Ixodes angiistus. ords of tlii.s tick biting humans in the USA and stated there was no suggestion of disease transmission in these instances.

Ixodes granulaius Supino (Fig. 82-85)

Ixodes gramddtus Supino, 1897:230-238; Asa- numa and Kosaka. 1955: 195-196; Keegan and Toshioka, 1957:19, PI. 19, 30; Arthur, 1957:683-694, Fig. 12-17, 19-24; Kawashima, 1963:103; Asanuma, 1965a: 11.3-120, 1965b: 398, Fig. 218.

Ixodes sp. 4, 51 and sp. 101 Asanuma and Seki-

kawa, 1952:107-116, Fig. 1, 2, 4, 6, 8, 1953:

99-112, Fig. 9, 10, 1.5, 16\ 17.

Discussion: Asanuma and Sekikawa (1952, 1953) reported the occurrence of three unnamed ticks of the genus Ixodes, including larval and nymphal fonns on rodents on Honshu, Kyushu, Hokkaido, and Shikoku. These unnamed ticks were tentatively numbered as Ixodes sp. 4 for larvae, Ixodes sp. 51 for nymphs, and Ixodes sp. 101 for adults. Keegan and Toshioka ( 1957) stated that Asanuma (1956a) indicated that these speci- Bhigham Young University Science Bulletin 120

S.SHIBATA

KiG. 82. Ixodes- finmuhitiis. female Biological Series, Vol. 15, No. 1 Ticks 121

Fic. 8.3. Ixodes nramdattis, male. Young University Science Bulletin 122 Bricham

Fig. 84. Ixodes g,r(inulnlits. nvmpli Biological Series, Vol. 15, No. 1 Ticks 123

Fig. 85. Ixodes granulatus, larva. 124 Bhicham Young University Science Bulletin

Biology: divergent; palps broad; the suture between

The following life-cycle data is taken from articles II and III sometimes indistinct; cornu;i Nadchatram (1960:220). absent; auriculae ;il)sent. In the female the scutum is cordate, longer than wide, widest an- The life cycle of the progeny of 3 female terior to the middle; hypostome blunt at tip with Ixodes graiuthitiis 6-7 distinct lateral denticles. In the male the reared in the laboratory on white rats palps are broiid ;iiid oval; the hypostome is distinctly notched ;it the tip uith l;iint Period indications ot scalelike denticles. in Days Females detached from host DiSTIUBUTION AND HoSTS: Females oviposited after 12-14 Arthur (1963) states tliat this species has Females continued oviposition for 16-20 ;in extensive distribution, including Britain, Ger- Larvae hatched out after 28-32 many (as two .subspecies), France, Kazak SSR, Larvae were fed after 15-25 Guryev Province. Arthur suggests this tick may Larvae fed for 4- 6 occur wherever sand martins nest. Hosts re- Engorged larvae quiescent after 6- 8 corded are swallows, Riparia sp., and the Japa- Nymphs emerged after 14-18 nese house nuutin, Delichon urbica dasijpus. Nymphs were fed after 14-42 Nymphs fed for 6-10 Engorged nvmphs (juiescent after 5- 7 Ixodes Itvidiis Adults emerged after 16-20 Adults fed after 28-56 ^ Collected and examined Adults fed for 10-14 O Collection records from literature above Total duration to complete life cycle 174-272 ^ Combination ot

Disease Relationship: Unknown.

Ixodes lividtis C. L. Koch (Fig. 86-89)

Ixodes lividus C. L. Koch, 1844:234; Ono, 1967: 217; Uchikawa and Sato, 1969:95-97.

Discussion: This tick had not been reported from Japan until recently, when Ono (1967) reported it from the nest of the swallow, Riparia riporia ijimae, from Hokkaido, and Uchikawa and Sato

(1969) collected it from the nest of Delichon urbica dasijpus in Nagano Prefecture. Three other lots of specimens were collected bv 406th Medical Laboratory personnel in Shimane and ( aimma prefectures. Ardiur (1963) discussed synonymy and distribution, and he considers 7. pUnnheus, I. cani-

•vugfl, and two subspecies, I. I. havaricus and /. /. ohotrUicus, both described by Schulze and M.\i> 26. Known distrihiitidn of Ixodes lividus. Schlottke, to be synonyms of lividus.

DlACXOSIS: BlOLOOY: j This is the only ixodid tick found on swal- Little is known of tlie biology of this species. lows in Japan, and while it h;is no unusual mor- .\rtliur ( 1963) st;ites that the males do not occur phological characters, the following points mav on the host, but their presence in the nests sug- serve to distinguish it from other species: body gests that copulation occurs off the host. In oval; the coxae have no internal spurs; in front Russia various biological aspects of this species j of the anus the anal groove is round; the have been studied by Glashchinska\a-Babenko posterior arms are almost parallel or slightly (1956) (see Babcnko', 1956). Xuttall and War- Biological Series, Vol. 15, No. 1 Ticks 125

r 126 Brigham Young University Science Bulletin

sonobe

Fig. 87. Ixiidcw tiiidus, male Biological Series, Vol. 15, No. 1 Ticks 127

Fic. 88. Ixodes liiidus, nymph. Bhicham Young University Science Bulletin 128

0,1mm -TTA-vfJ-o,

Kic;. 89. Ixodes litUlus. l,ir\a. BioLOciCAL Sehies, Vol. 15, No. 1 Ticks 129 burtons (1911) comineiits on the biology ot ,' Ixodes monospinosus . /. canhu^ii apply to /. Uvidus. In the 406th •0 e Medical Laboratory eight engorged females laid 257-810 (average 5.58) eggs,"which hatched 25- 28 days after the beginning of oviposition. Larvae were placed on rabbits" ears, and 13-17 days later they molted to nymphs.

Disease Relationship: UnknowTi.

Ixodes monospinosus Saito (Fig. 90)

Ixodes monosiiinostis Saito, 1967:107-112, Fig. 2-9.

Discussion:

Saito's ( 1967 ) description was based on a single female removed from a mountain climber in Niigata Prefecture. Saito illustrated (Fig. 2-9) the dorsal aspect, capituluni, genital and anal regions, spiracular plate, tarsi and coxae (palpi may have been missing). The holotype was not available for study, but the following diagnosis was provided by Saito.

Diagnosis:

The female of /. monospinosus is easily differentiated from known ixodids from Japan and other countries bv the following series of features: hvpostome long, with four dental files on each side; auriculae poorly developed; coxa

I with onlv a spinelike, stout, inner spur, and the posterior margins of coxa I are thickened transversely; coxae II-IV each have only an external spur and a moderately convex ventral surface; scutum and carinae well defined, lateral grooves distinct, large punctations on median- posterior areas, and posterolateral margins slight- Iv concave.

Distribution and Hosts:

Saito lists man as the host and the specific locality as Mt. lide, Mitakambara-gun, Niigata Prefecture, Japan. A single female specimen collected by the flagging method in Mie Pre- fecture was loaned through the courtesy of Dr. Kitaoka and illustrated in this paper.

Biology: UnknowTi.

Diskase Rel.\tionship: Unknown.

Ixodes nipponensis Kitaoka and Saito (Fig. 91-94)

Ixodes nipponensis Kitaoka and Saito, 1967:74- 83, Fig. 1-19. University Science Bulletin 130 Brigham Young

I.O m m 0.5 m m

Kic. 9(). Ixodes monospitwsus, feni;de. Biological Series, Vol. 15, No. 1 Ticks 131

^,i}hTauJ(i

Fic. 91. Ixodes iiippuriensis, fem;Ue. BiiiGHAM Young University Science Bulletin 132

'*^ <^^«i<.»«^<»-

Fit:. 92. Ixodes iii/j/JDiicii.vi.s, male 133 Bi()loc:k;al Series, Vol. 15, No. 1 Ticks

Fio. 93. Ixodes ni/j/ioiicruvw, nvmpli. Bhigham Young University Science Bulletin 134

Fic;. 94. Ixodes nipponcnfii.s, huM Biological Skries, Vol. 15, No. 1 Ticks 135 mature tick.s, Ixodes .sp. LA (larva) and Ixodes badger, hare, weasel, and man. Immature stages sp. NA (nymph), reared from egg.s laid by an are found on small rodents and birds. In a forth- undetermined .species of Ixodes. These immature coming publication by Kitaoka detailed informa- ticks were similar to Ixodes sp. 2 and Ixodes sp. tion will be given. 54 of Asanuma and Sekikawa (1952, 1953), hut separable from these by a combination of sev- Biology: eral morphological characters. 1962a ) be- Ono ( An engorged female (67-J-0341) laid eggs in lieved that Ixodes sp. 2 and sp. 54 were the im- the 406th laboratory and the progeny were used mature fonns of /. persidcatus and also expressed for life cycle studies. doubt about the identitv of /. /). perstdcatus, which had been used bv Saito, Ohara, and Una- gami (1960).

Kitaoka and Saito ( 1967 ) showed that Ixodes sp. LA and Ixodes sp. NA were larval and inmphal fonns of Ixodes nip])onensis. Though the adults of nipponensis are very similar to those of persidcatus, the immature forms are separable from each other. Consequently, records of ricinus reported bv earlier |apanese workers might possiblv include nipponensis. The material recorded in Keegan and Toshi- oka (1957) included onlv persidcatus, but these specimens were reexamined bv Yamaguti with the help of Dr. Kitaoka, and they concluded that several lots previouslv identified as persidcatus actuallv included nipponensis. Kitaoka pointed out that the specimen on which the illustration of persidcatus persidcatus (PI. 34) in Keegan and Toshioka ( 1957 ) is based, is nipponensis.

DiAGXOSis:

This species is quite similar to /. persidcatus but mav be distinguished from that species in that the bodv is smaller and is covered a little more denselv with thick setae; coxa I has a slightlv shorter spur than persidcatus: in the male, the internal spur on coxa I is shorter and reaches the anterior margin of coxa II, while it is long and partially covers coxa II in perstdcatus; the spiracular plate is much elongated longitu- M.\p 28. Known dLstrihution of Ixodes nipjioneruis. dinallv and ellipsoidal; in the female, the internal Disease Relationship: spur on coxa I is shorter and does not cover K of coxa II; the marginal dorsal setae are forked, Unknown. thicker and approximately 1.7 times longer than Ixodes ovatus Neumann the pointed, simple setae in per.sulcatus. More- (Fig. 95-98) over, Kitaoka and Saito (1967) and Ono (1962a, Ixodes ovatus Neumann, 1899:116-118, Fig. 2, 3 1966) compared the differences between im- (in part), 1904:452-453, 1911:18; Donitz, mature stages and concluded thev were on a 1905:132; Kishida, 1930a:3; Nakamura and specific level. See also diagnosis of per.sulcatus. Yajima, 1937:141; Keegan and Toshioka, Distribution axd Hosts: 1957:29-30; Morel, 1963:925-928. Ixodes ricinus var. ovatus: Nuttall and Warbur- /. nipponensis has been recorded by Kitaoka and Saito (1967) from Honshu, Kyushu, Shiko- ton, 1911:158. ku, and Sado Island, and from a few localities in Ixodes japonensis Neumann, 1904:458-459, 1911:

Hokkaido by Ono ( 1966). Records of the 406th 22; Nuttall and Warburton, 1911:208-209, Medical Laboratory include Shiga, Shimane, Mie, Fig. 200, 201; Kishida, 1930a: 3; Nakamura Hiroshima, Tokyo, Kagoshima, and Kochi pre- and Yajima, 1937:145-146, PI. Ill; A.sanuma, fectures. Hosts recorded are cattle, horse, dog, 1947b: 971, Fig. 2765; Keegan and Toshioka, 136 BmcHAM Young University Science Bulletin

Laboratory life cvcle of Ixodes nipponensis (reared on rabbits) Generation and Stai^c Phase Period in Days Remarks

F, Larva Feeding 5 Larvae were placed on host on F, Larva I'ostpaiasitic 30 5 July 1967. (Preniolting) F, Nynipli Feeding 4 F, Nymph Postparasitic 33 Only 2 nymphs molted to (Preniolting) females.

The rearing experiment extended from July 1967 to October and the air-conditioned room temperature did not exceed 25 C.

1957:20-21, PI. 31, 32; Saito, 1959b 199-203, described ovatus based on two females from a 1960a; 303-321, 1962a: 127-146, 1962b 147-159, horse and a dog at Saga (No. 704 and 705) as 1964:59-66; Saito and Ohara, 1961:l-.32; Saito, separate from the other t\\'o. Nuttall and War-

Ohara, and Unagami. 1960:323-329; Saito burton ( 191 1 ) cited Neumann's description

et al„ 1965:143-159; Asanuma, 1956a:91-96, ( 1904 ) , and in ;iccordaiice with Donitz's opinion 1957:2.34, 1965a:] 13-1 16, Fig. 7, S, 1965b: they noted that the ditferences between the two 398, Fig. 217; Ono, 1966:62-68. species were only varietal. It is apparent that Ixodes frecjttens Ogura and Takada, 1927:201- Nuttall and Warburton did not see the type 202, PI. Ill; Schulze, 19.30:294-303, Fig. 10; specimens. Therefore, they placed ovatus as a Kishida, 1930a:3; Nakamura and Yajima, variety of ricinus and treated jaj)onensis as a 1937:146-148, PI. IV; Yajima, 1942:507-509, valid species separate from ovatus.

Fig. 6; Itagaki, Noda, and Yainaguchi, 1944: Neumann's original description of /. jajioncn-

1-97, PI. 2, 1959:1-118. PI. 2. SK ( 1904 ) did not include illustrations. Later, Ixodes carinatus Kishida 1930a: 4-5; Nakamura Nuttall and Warburton (1911) provided illus- and Yajima, 19.37:140; Keegan and Toshioka, trations (their Fig. 200 and 201) and a rede- 1957:26, PI. 41. scription of jajwnen.sis based on Neumann's type specimen collected in the vicinity of Tokyo. Ixodes sp. 6 and sp. .52 Asanuma and Sekikavva,

1952:107-116, Fig. .3, 9, 1953:99-112, Fig. 11, Pomerantzev ( 1950 ) considered ovatus as a 12; Asanuma, 19.55:1240-1242; Ono, 1962a:24- valid species occurring in Japan, but this appears 29, Fig. L5, 1962b; 155, 1966:62-68. to be merely a citation from Neumann's original ?Ixodes shinchikuensis Sugiinoto, 1937:330-331, record.

PI. VI and IX. Ogura and Takada (1927) described /. fre- ?Ixodes taiwanetiiis Sugimoto, 19.36:336-.346, PI. quem\ but Keegan and Toshioka (19.57) sug-

II, 1937a :331-333, PI. VII and VIII. gested, "After a study of hosts and localities, it

seems clear that frequens is a synonym" (of Discussion: jafwuensis-ovatus). Thev also examined the

In the Japanese literature this tick has been holotvpe specimen of /. carinatus Kishida, 19.30 reported as /. joponensis, I. frecpicns. and some- and stated that "the holotype is very much like times /. carinatus. an example of japonensis, although the basis Ixodes ovatus was originally described bv capituli mav be relatively wider than in that Neumann (1899) on the basis of material from species." More recently Morel (1963) showed

Amur and Japan. The Japanese material includ- that /. jajwnrnsi.s: I. fre(juens, I. carinatus. I. ed four female specimens, one from a horse at shincliikuensis, and /. taixcanensis were all syno-

Akita collected by Taguchi (No. 702), the others nyms of /. ovatus. The latter two species were from a Lepus sp., a horse, and a dog at Saga described bv Sugimoto on the basis of Taiwan collected by Yainaguchi (No. 703, 704, 705 re- specimens, ;ind Morel (1963) expressed uncer- spectively, in the Neumann Collection in the tainty regarding their status, probably because School of Veterinary Medicine in Toulouse). of lack of specimens. Although the holotype According to NuttaJl and Warburton (1911), specimen of carinatus was not available for this these Japanese specimens contained typical study, the original illustration by Kishida ricimis," and subsetiuently Neumann (1904) re- (1930a) fits the characteristics of ovaltis and

'Nlorvl llinl (196}) slimvcd tlicsc riciiius I No r02 iind 703) wcro iidiwllv /. pcrsulcnlin. I>nl llicsi' uiiKlU or ]iii(;lu mil iiuUiilo / ponrnsii Kitnnkn and Saito. Biological Series, Vol. 15, No. 1 Ticks 137

•^ -;£^^-i*t-<5-c^

Fig. 95. Ixodes ov(du.s, female. Bkioham Young University Science Bulletin 138

/O' ^-^^r^^Otf^o^

Fit:. 96. Ixodes onitim. male Biological Series, Vol. 15, No. 1 Ticks 139

'/^^A<

Fig. 97. Ixodes ovatu.s-, iijmph. 140 Bhicham Young University Science Bulletin

0.1 m m

BuucKi ;

Biological Slries, V'ol. 15, No. 1 Ticks 141

supports Morel's opinion. The only report of this species as ovatiis was Ixodes ovatus given by Kishida (1930a), but he also reported ja))onensis on the same page. Asanuma and ^ Collected and examined Sekikawa (1952, 1953) reported and described O Collection records from literature larval and nyniphal ticks parasitic on rodents, giving them temporarv numbers such as Ixodes sp. 6 or Ixodes sp. 52, because the immature stages had not been properly associated with ^nr Shirrbeshifc^fc^O/Kushir adult fonns. Later Asanuma (1957) showed Is / -^vTlburi^idaka Oshima I that these numbered ticks, Ixodes sp. 6 and Ixodes sp. 52, were larval and nvmphal forms of Ixodes japonensis (= /. ovatus) respectively. Ono (1962a, 1962b. 1966) reported immature ticks from rodents in Hokkaido and gave them the numerical designations used by Asanuma and Sekikawa. CigataOr OFukushima C'umma ToyamaQ^-r- .^_u.„.Tochigi DiAGXOSIS: FukuiQj ^^»"°, Tokyo ^ . The dorsal aspect of the male of this species A°jrK'KvSto^nianash Chiba is similar to /. nipponensis and /. perstdcatus, Shizuoka -' Kanagawa ^ 'Okayama"^'^ Hyogo but from a ventral view it may be easilv distinguished from those two species in that coxae II

and III are lacking spurs, and the posterior jKumamoto halves are covered with milky white, membra- Kagoshima I

nous, eavelike elevations ( = carinatus Kishida, 1930), and consequentlv the posterior margins are smoothly convex; coxa I has a short, blunt

internal spur; the spiracular plate is elongate, the L/W ratio is more than 2.0 (approximately 2.3); Map 29. Kno di.stribiition of Ixodes ovatus. the dentition of the hvpostome occurs on the

apical half, the posterior end is on the same preparasitic larval period was about 40 days. level as the suture between palpal articles II and Feeding time of larvae varied according to tem- III. In the female, the basis capituli has well- perature and was from 2-6 days in summer and defined cornua and round porous areas; coxa I from 7-10 days in late autumn on Angora rab- has a short, blunt internal spur; on coxa II, the bits, and from 4-7 days on laboratory mice. The

spur is absent and the posterior margin is convex postparasitic period was 24-.32 days at 30 C, and with the elevation similar to that of the male. newly hatched nymphs attached to the host 14 days later. Nymphs fed on the host for about DlSTRIBlTIOX .\ND HoSTS the same period as, or one or two days longer

Morel ( 1963) states that this species occurs than, larvae, and they molted into adults 30-.37 in oriental regions, including China, Formosa, days after they dropped off the host. The pre- Bunna, Nepal, Sikkim, and Japan, and that hosts parasitic period for adults was 20-30 days, and of the adults arc large mammals. .'Kccording to the unfed males began mating immediately after Asanuma (1965a), adults are found on hares in they were put together with females on the host. great numbers, especially in spring, but imma- Females fed on the host for 9-13 days in spring ture forms are not found on hares. Asanuma and autumn, 7-10 days in summer, and 12-16 and Sekikawa (1953) and Saito (1959b) state days in winter. The engorged females laid eggs that immature fomis are found on small rodents. in .5-7 days in summer and in 10- L5 days in Detailed infonnation on the distribution and spring, and the oviposition period varied from hosts of this species in Japan is given in 10-40 days, depending on the room temperature. Appendix 2. Saito also observed the behavior of mating and oviposition. He stated that summer seemed to Biology: be an unfavorable time for feeding of adults in Saito (1960a) studied the life cycle of the laboratory. Ixodes ovatus (as /. japonetviis) with laboratory animals as hosts (Angora rabbits and mice). Disease Relationship: Eggs hatched in .32-38 days at .30 C, and the The relationship of this species to human dis- )

142 BiiiciiAM VouNC Univebsitv Science Bulletin

ease has not been established. Saito ( 19621) occurred in Japan and suggested that there mav reported the result of experimental transmission be two or three separate species contained in the of Piisteurclld lularen.sis with /. idjxmciisi.s. The Japanese population of the ricinus group. Asa- adults of 7. japonensus ( = /. ovatus), detached numa and Sekikawa ( 1952, 19.53) described im- from hares dead of tularemia, were found to be mature ticks parasitic on small rodents and gave infected and shown to be capable of transmitting them temporary numbers, i.e., Ixodes sp. 2 for P. tularcn.si.s to Angora rabbits. Attempt.s to kirvae and Ixodes sp. .54 for nymphs. They demonstrate transovarian transmission were neg- pointed out that these numbered ticks were ative. very similar to immature forms of ricinus, but avoided a definitive assignment. Ono (1962a) Ixodes persulcatus Schulze examined a large number of reared larvae of (Fig. 99-102) Ixodes persulcatus and considered that Ixodes Ixodes persulcatus Schulze, 1930:294-303, Fig. sp. 2 and sp. 54 reported by Asanuma and Seki- 3, 5, 6, 8, 9. Morel, 1963:925-928"; Ono, 1966: kawa were the immature forms of Ixodes per-

62-68, Fig. 1; Kitaoka and Saito, 1967:82. sulcatus. Ixodes perstdcdtu.^ i)ersidcatus Schulze, 1930: Keegan and Toshioka (19.57) state that 294-303, Fig. 3, 5, 6; Asanuma, 1947b :971, Ixodes ricinus var. niiiiazakicnsis Sugimoto, 19.37 Fig. 2764, 1965a: 113-116, I965b:397, Fig. 215; appears to be persulcatus persulcatus. But in the Keegan and Toshioka, 1957:21-22 (in part). light of current knowledge, the written descrip- Ixodes sp. 2 and sp. 54 Asanmiia and Sekikawa, tion and the illustrations are too poor for taxo- 1952:107-116, 1953:99-112, Fig. 13 and 14; noinic discrimination, and this material could be Asanuma, 1955:1240-1242; Ono, 1962a:24-29, l)er.sidcatus or nipponensis. Since the adults of

Fig. 4-6, 8, 10, 12, 1,3, 1962b: 155, 1966:62-68, persulcatus are very close to nipponensis and the Fig. 1. specimens reported as ricinus by earlier Japa- ?Ixodes ricinus: Neumann, 1899:166 (in part), nese workers are difficult to obtain, there are

1904:452, 1911:12-13; Nuttall and Warbur- only a few reliable collection records of /. per- ton, 1911:1.56, 1915:433; Ogura and Takada, sulcatus. 1927:199-201, PI. XI; Kishida, 1930a:2, 1936: Diagnosis: 142; Nakamura and Yajima, 1937:142-145, PI. species is closest to /. nipponensis in the 11; Yajima, 1942:504-508, Fig. 5; Itagaki, This fauna and also to ricinus in other Noda, and Yamaguchi. 1944:1-97, PI. 1, 1959: Japanese regions. Kitaoka and Saito gave a critical 1-118, PI. 1. ( 1967) ?Ixodes persulcatus ])ers-idcatus: Saito, 1958:70, comparison of these three species and indicated 1959b: 199-20,3, 1960a:.303-321, 1962a: 127-146, that adults of persulcatus may be distinguished 1962b: 147-149; Saito and Ohara, 1961:l-.32; from nij)j)onensis bv the larger bodv size as well Saito, Ohara, and Unagami, 1960:32.3-329. as by a combin;ition of characters used in the ?Ixodes ricinus var. miyazakiensis Sugimoto, key. In addition, the following characters of stages given Kitaoka and Saito 1937:603-604, PI. 1. immature bv (1967) and Ono (1962a, i962b, 1966) are u.se- Discussion: ful in separating the three species. In the This tick had been widely reported in the larvae, the scutal setae are of medium length, Japanese literature as Ixodes ricinus ( Linnaeus, while they are apparently shorter th;m the post- 1785). Schulze (19.30), in his original descrip- scutal setae in both nipjwnensis and ricinus; the tion of /. persidcatus, examined two female speci- livpostomc of the nvmph is blunt at the tip mens from that speci- Hokkaido and stated these rather than pointed as in nipjwnensis: the post- mens appeared to be conspeeifie with a small scutal setae are not forked; the auriculae and Amur specimen ( one of the type scries ) show- spurs on the coxae are more distinct in persul- ing all the characteristics /. persidcatus. of He catus than in nipponensis both in the larval also stated that specimens described bv Ogura and the nvmph;il stages. See diagnosis of nip- ;uid ;is Takada (1927) ricinus from Hokkaido jionensis. apparently belonged to his new subspecies, I. p. persulcatus. Keegan and Toshioka (19.57) re- Distribution and Hosts: ported that all the material available at that time The tvpe material was collected at .\inur.

was reexamined and identified ;is Ixodes persid- Schulze mentions other material from Hokkaido

catus j>rrstdcatus. Since then, the ]apanese popu- which had been described ;is ricinus by Ogura lation had been thought to include onl\' junstd- and Takadii (1927). According to Pomerantzev catus persulcatus. Later Asanuma (1 96.5a) ex- (19.50), the distribution of this .species is from pressed doubt that onlv persidcatus persidcatus the Pacific Ocean to the Furopean p;irt of the .

Biological Series, Vol. 15, No. 1 Ticks 143

/ okCXuiA,

Fig. 99. Ixodes persulcatu.s, It'inalf 144 Bricham Younc University Science Bulletin

^.oAv^ujd^

Ki<;. 1(K). Ixodes persulcatus, male. 145 BioLocicAi. Sehies, Vol. 15, No. 1 Ticks

-^. 'TriAjiu'Ju-

Fic. 101. Ixodes persulcatus, nymph. 146 Bricham Younc; IInivehsitv Science Bulletin

j^m^Ti/)

Fit:. 102. Ixodes persulcntus. Iar\; .

Biological Series, Vol. 15, No. I Ticks 147

USSR, and al.so in northern Germany. The dis- studied by Russian workers who were concerned tribution in Japan i.s given in Appendix 2. The with the transmission of spring-summer encepha- only Korean record is from a horse at Kankyo- litis to man. The life cycle was studied in nature lioku, now known as Hamgvong-pukto, by Ita- by Serdukova, and the infomiation is given by gaki et al. (1944, 1959). Kishida (1936) men- Pomerantzev ( 1950 ) . The length of the develop- tions that /. ricintis was recorded from North mental cycle depends up(m environmental fac- Korea bv Akasawa in 192S. We have been un- tors and is said to take three years in the boreal able to confinn this record or to find any paper taiga in the USSR in warmer regions; and, in by Akasawa written in 1928. Asanuma and the presence of abundant host animals for all the Sekikawa (1953) listed the hosts of ricimis and three active stages, the tick develops in a re- j)ersulcatus\ citing the records by Kishida duced two-year cycle. Ixodes j)ersulc(itus is a

) in ( 193()a ) and Nakamura and Yajima ( 1937 typical forest inhabitant and is most abundant They are human, cow, horse, wild rabbit, deer, mixed deciduous-coniferous forests. dog, fox. leopard, weasel, martin, hedgehog, The seasonal occurrence of persulcatus in sheep, goat, and rat. Pomerantzev (1950) stated Niigata Prefecture, Japan, was studied by Saito that the main hosts of adults in the USSR are (1959b) through collections from wild animals wild ungulates such as deer, elk, roebuck, and or from vegetation using the flagging method. the Siberian stag. Hosts of larvae and nymphs The following biological information was iiuoted are small mammals ( Rodentia, Insectivora, Car- from Saito's extensive studies, but as noted in nivora) and birds. Nymphs are also parasitic the discussion, this might be applicable to per- on the Amur hedgehog, dormouse, scjuirrel, sulcatus. provided Saito's material definitely did hare, and wood-hen. Both adults and nymphs not include nipponensls. Both immatures and attack man. adults were found together on hares. The adults were generally found active on the host through- hodes persulcanjs and/ nanus out the year, showing a slight increase in num- examined bers in spring and late autumn. Both nymphs 1 ^ Collected and abundant on the host from 2 Q Collection records from and larvae were most literature April through July. Larvae were not collected Combination of 1 and 2 3 3 on hares in winter and midsummer. The imma- 4 A Collection records of ture fomis were also rare on Apodemtis specio- / ncinus stis sadoensis in summer. Using the flagging 5 <$ Combination of 2 and 4 " StiiribestiiftnCy''i^'>'^"*"° method, all stages were collected from vegeta- 6 Combination of 1, 2 and 4 O ^^ThIa^,-^'T°''0':l'i tion, and the peak abundance was from summer OshimaT^ to early autumn. Saito ( 1960a ) studied the laboratory life cycle and mating behavior, using Aomofi chiefly the Angora rabbit as the host animal.

•/ ®'wate I Females fed for 13-18 days in spring and autumn Yama9ata( 7-14 in summer. He found that if fe- kMiyagi and days SadoC^ 1} males leave the host before becoming fully en- they ^Niigata^-r *A Fukusfiima gorged they weigh less than 70 mg, and rarely feed on another host. Females laid eggs J.Saitama -O— ^ Naganoflt)^ in summer and 9-14 Kyoto . ? "JrllS'^"tV\ fQ.^® spring. Oviposition lasted 10-14 days un- Shimanejn^Hyogd^t^_X '/K'iO' ' days in Karwgawa Yamanaslii , conditions, and larvae " der various environmental :Mie"~^ / hatched out in 30-36 days at 30 C. For larvae ,Ehime5%'f*\-^^ the preparasitic period was approximately 30 Kuniamoto days, and they fed for periods of from 3-13 days. after larvae had Miyazaki Nymphs emerged in 17-19 days dropped off the host. Ten days later they fed on the host for periods which varied from 4-15 days. Adults emerged in 25-30 days at 30 C. The minimum time for the total life cycle in the laboratory was approximately 4.5 months. Map .30. Known distriUntion of /ayx/cs prrsulrcilus and behavior of /. ricniuv. An interesting peculiarity of the persulcatus was observed by Balashov (1954). Biology: The dropping of engorged female ticks from the coincides with the time when the cows were The biology of /. pcrsulcutm has been host 148 BiuGHAM Young University Science Bulletin in the pasture. If cows were pastured during Diagnosis: tile davtinie, tlien tlie ticks dropped during the We (juote directly from Keirans and Kohls daytime, hut if cows were pastured during the (1970): "Ixodes ))hilipi n. sp. superficially re- night, ticks night. then the dropped at sembles /. kohlsi described by Arthur (1955) from Eudtjpttda minor or from its nest, and keys

Disease Relationship: to /. kohlsi in Roberts (1960), However, /. Ixodes persulcaius transmits the virus of philipi is much more hirsute dorsally and has a spring-summer encephalitis to man in the USSR. less elongate scutum. Ventrallv /. ))hilipi has a Ponierantzev (1950) states that the causal smaller, less distinct external spur on coxa IV, agent ot cattle piroplasniosis can he transmitted and spur on trochanter I\' obsolete." experimentally. Saito ( 1962b) studied the trans- Of the Japanese Hcks associated with sea mission of PasteurelUi tularensis by /. perstilca- birds I. philipi most closely resembles /. signatus tus and found that adult female specimens of and kevs out to that species. However, females persulcatus, collected from wild hares which had may be distinguished from /. sigmitus by the died of tularemia, were infected with P. tularen- following diagnostic characters: Dorsum pos- sis and that nymphs and larvae that were ex- terior to scutum uniformly covered with white perimentally infected with the causal agent of hairs; coxae I-III anned with a more pronounced tularemia transmitted it from host to host. He external spur; anal groove markediv constricted also demonstrated transstadial transmission, both and subparallcl posterior to anus giving "key- from larvae to nymphs and from nymphs to hole" shape; basis capituli ventrally with broadly adults, but he was unable to demonstrate trans- rounded posterior margin, with straight lateral ovarian transmission. The role of /. persulcatus margin anterior to auriculae; auriculae broadly as a vector of P. tukirensis from hares to man triangular or nearlv right-angled ridges; porous has not been established. areas well separated by about half the width of one of them; hypostome untoothed at base, about Ixodes philipi Keirans and Kohls (Fig. 103) Ixodes philipi Keirans and Kohls, 1970:725-72e, Fig. 1-8.

Dl.SCUSSION:

The description of this species was based on two partially replete females collected by H. E. McClure in 1951 at Sanganjima, Iwate Prefec- ture, Japan. Keirans and Kohls (1970) (juoted from a personal communication from Dr. Mc- Clure in which he described the collecting area ". as . . high islands of volcanic ash covered with pin(> and deciduous forests. In the loose soil beneath the trees the Streaked Sheanvater, Pttffimis leucomelas, and the Madeiran Storm Petrel, Oceanodroma castro, dig nesting tunnels about arm-length deep." Ticks were collected during an avian blood survev but it isn't certain which of the two species of birds was the host from which the ticks were collected. Japanese ornithologists now refer to the Streaked Shear- waster as Calonectris leucomelas and the Ma- deiran Storm Petrel is sometimes called the Ma- deiran Fork-tailed Petrel. Keirans and Kohls (1970) indicated that the female characters ful- fill the criteria for placement of /. philipi in the subgenus Scaphixodes Schul/.e, 1941 but because of the lack of males and immature fomis thev are hesitant to do so. More extensive tick collections from sea birds are essential for ecological and taxonomic studies. M.AP 31. KniiNMi ilistrihiitioii i>t Ixo(lc.\ philipi. Biological Series, Vol. 15, No. 1 TlCK.S 149

10.3. Fig. Ixodes philijn, female. ( .After Kierans and Kohls 1970) 150 BHif;iiAM VouNG University Science Bulletin one fourth as long as the wliolc length, with saliences present in the female; cornua lack- more prominent teeth on inner file. ing; palp article III slightly longer than Article II, both articles II and III relatively Distribution and Hosts: shorter and broader than in the female. Known only from Ptiffiniis Icucomchis (Tem- liijpostomc about three-fourths as long as minch, 1S35) or Oceanodwiiia castro (Harcourt, palpi, relatively shorter than in female; den- 1851), Sanganjima Island, Iwate Prefecture, ticles indistinct. Scutum: Cervical grooves Japan. distinct, nearlv straight, diverging; two longitudinal, Biology; nearly straight, slightly diverging grooves posterior to cervical grooves. Unknown. Between these is a single, indistinct, longitudinal groove with faint cross-bars which Disease Relationship : extend the Unknown. to paired longitudinal grooves. A dorso-lateral sclerotized patch on either Ixodes signatiis Binila side of the body at a level between coxae (Fig. 104-107) II and III. Posterior margin of scutum ir- regular in outline, and tenninating in a Ixodes signatus Rirula, 1895:.357-358, PI. 1, Fig. point in most specimens examined. The 10-13; Nuttali 1911:261-264; and Warburton, scutimi does not cover the entire dorsal sur- Kishida, 19.30a:2; Znmpt, 1952:18; Asanuma, face within the lateral groove. This is par- Okubo, and Fnkuda, 1955:85-86; Asanuma ticularly true on the posterior fourth of the and Fukuda, 1957:147-159; Keegan and To- bodv. Palpi more noticeably haired than in shioka, .35 1957:22, PI. and 36; Asanuma, the female; especially distinctive are a 1965a: 11.3-116, Fig. 216. 1965b:398, group of short, stout setae on the lateral, Ixodes parvirostris Neumiinn, 1901:284. dorsal aspect of articles II and III; terminal Ixodes eudijptidis var. signata: Neumann, 1904: setae also are more conspicuous. Legs and 451. coxae as in female. Coxal spurs on legs III Ixodes eudi/ptidis signatus: Neumann, 1911:21. and IV may be relatively larger than in female. Spiracular plate oval rather than Discussion: circular; macula displaced anteriorly; gob- This tick was first reported from Japan by lets as in female. Genital aperture between II. Genital start Neumann ( 1901 ) as Ixodes parvirostris, which coxae grooves to converge he described from eight female specimens taken at level of anterior margin of anal groove. from the connorant, PIxdacrocorax pelagicus, on Median and adanal plates not distinct; a

Yezo, Japan. Yezo is the archaic term for Hok- slight trace of sclerotization and pigmenta- kaido. Nuttali and Warburton made parviros- tion in the areas usually occupied by the tris a synonym of sigiiatus in 1911. Kishida adanal shields, and a more extensive pig- (1930a) and Znmpt (1952) cited Neumann's mented area in the usual location of the collection record, but gave no new information. ventral shield; this pigmented area has ir- Asanuma, Okubo, and Fukuda (1955) reported regular lateral and posterior margins. Aiud an infestation of the black-tailed gull. Lams groove rounded anterior to anus; the arms crassirostris, by this tick at Kabujima, Aomori of the groove are curved throughout, and Prefecture, Honshu. Additional specimens from converge posterior to the anus; distance be- the same locality were collected by 406th Medi- tween these arms is greatest a short dis- cal Laboratory personnel. tance posterior to the anus.

Diagnosis: Distribution and Hosts: This species is distinct from other Ixodes sp. Znmpt (1952) has reviewed the distribution and may be easily identified using the key. of signatus and other species of ticks infesting Description of the male by Keegan and Toshi- sea birds. This species has been collected on oka (1957) is based on material collected from only a few occasions in other areas. It has been Lariis crassirostris at Kabujima, Japan: found on the connorant on Hokkaido, Japan, Body widens abruptlv from scapular the Aleutian Islands, and from t\vo locations in region; with nearly parallel lateral margins; California. It was taken once from a seal in the posterior body margin bluntly rounded; lat- Aleutian Islands and on several occasions from eral groove distinct, (^apituhii)i relatively the black-tailed gull in its nesting areas at Kabu- short and broad, but without the lateral jima, Aomori Prefecture, Japan. Biological Series, Vol. 15, No. 1 Ticks 151

i JHIBATA

Fig. 1()4. Ixodea signatus, female. BiiiGHAM VouNC University Science Bulletin 152

S.SHIBATA

Fig. 105. Ixodes ni^iuittia, male. Biological Series, Vol. 15, No. 1 Ticks 153

RuucKi

Fic. 106. Ixodes signatus, nymph. Univebsity Science Bulletin 1.54 Bhigham Young

W t'ia'iin\un\ii.

Fio. 107. Ixodes sisiuitiis, larva. Biological Series, Vol. 15, No. 1 Ticks 155

Ixodes signaws 156 Bricham Young University Science Bulletin

RTSUTSUMI

l-'ic. 1()8. Ixodes ximplcx Kimplrx, female. Biological Series, Vol. 15, No. 1 Ticks 157

Fic;. 1()9. Ixodes .simplex simplex, nymph. 158 BiuGHAM Young University Sc:ience Bulletin-

ferentiated from other Japanese ix(Klids by the

following characteristics: "In /. tanuki, with a

long sharp internal spur on coxa I, external spur

is absent on coxa I and II each. These morphological features are not observed in the other ticks. Ixodes ozarkus Cooley, 1944, of North

America, is thought to be resembling /. tanuki, but measurements of all parts of ozarkus are generally larger than those of tanuki. The for-

mer has an external spur on coxa I and II each, but these are lacking in the latter. The spiracu-

lar plate of ozarkus is elongated oval in shape,

but oval in /. tanuki."

Distribution and Hosts: Distribution and hosts of the species are at present limited to the badger in Niigata Prefec- ture, and there are no reports from other areas. Dogs have been suggested as possible hosts.

Ixodes tanuki . , ? 1

Map 33. Known distribution of Ixodes simplex simplex.

Biology :

Nothing is known concerning the biology. Males probably do not feed, or they may feed very rapidly and then quickly hide themselves.

Disease Relationship: Unknown.

Ixodes tamtki Saito (Fig. 110-111)

Ixodes tanuki Saito, 1964:59-66, Fig. 1-9.

Discussion: Ticks taken from a badger, Nyctereutes pro- cijonoides viverrinus, m Niigata Prefecture, Ja- pan, were described liy Saito (1964). He collected one male and three females in 1959 and an additional female in 1963. The specific name "tanuki" was derived from the Japanese common name for the host. Since the specimens were not

available for study, the following infonnation is taken from Saito's paper.

Diagnosis:

Saito states thai /. taiuiki mav be easilv dif- Biological Series, Vol. 15, No. 1 Ticks 159

1 .0 mm

0.2 mm 0.5 mm

Fic. 110. Ixodes tdiiuki. ffiiiale. 160 Bhicham Vounc University Science Bulletin

1 .0 mm

Fic;. 111. Ixodes tanuki, male. .

Biological Series, Vol. 15, No. 1 TiCK.s 161

Ixodes turdtts Nakatsiiji the Japanese specimen, the median posterior (Fig. 112-114)

Ixodes tardus Nakat.suji. 1942:292-294, Fig. 4; A.sanuma and Ko.saka, 1955: 192-194, Fig. 1-4; Asanuma and Nakagawa, 1955:549; Kee- gan and Toshioka, 1957^24, PI. .38; Asanuma, 1965a: 113-1 16.

Ixodes sp. 1 Asanuma and Sekikawa, 1952:107- 116. Discu.ssiON:

The description of /. tiinltis Nakatsuji, 1942 was based on seven females taken from the thrush, Turdus celacnopus ceJaetwpus, on Ha- chijo Island. .\dditionaI infomiation concerning distribution, hosts, and morphology of the species was given by Asanuma and Kosaka ( 1955 ) Diagnosis:

Asanuma and Kosaka ( 1955 ) commented on the close similarity between I. turdus and I. brunneus Koch. 1844 and were of the opinion that the following characters may serve to distinguish the two species: the scutum is large, long and rhombic, with a fairly distinct lateral angle at the widest part, the postlateral margin is slightly concave (distinct from brunneus in this); the auriculae are rectangular, well developed, and project externally ( the tip is visible from the dorsal side); the hypostome is 4/4; the anal groove converges posteriorly; the coxae M.\p 35. Known distribution of Ixodes turdus. have well-developed external spurs. area is flattened rather than convex, and Distribution and Hosts: the hairs are not quite as heavy as in This species has been collected only in Japan. brunneus. Quite likely this is Nakatsuji's Asanuma and Kosaka ( 1955 ) have collected (1942) species and if so the description is this species from a variety of birds, and on a few poor and the figures are inaccurate, espe- occasions from Apodemus speciosus. The majori- cially with regard to hypostome, auriculae, t\' of these collections were made in the Izu- and the comua. I should like to mention Schichito Islands ( Seven Islands of Izu ) soutli too that the 2 nymphs accompanying the fe- of Tokvo Bav, and other specimens were taken males are very similar to those of /. cali- in Chiba, .Shizuoka, and Tochigi prefectures, all famicus Banks, which is based on nymphs on Honshu. Hosts listed by them are given in only, and which is definitely brunneus .Appendix 2. (Cooley and Kohls, 1945:215).' Biology: Since validity of this taxon remains in (juestion,

Unknown. the name /. turdus used in this paper is tenta- Disease Relationship: tive. Unknown. Ixodes iiriac White White, 1852:10; Keegan and Toshi- Remarks: Ixodes uriae oka, 1957:.30; Asanuma 1961:181, 1965a: 116. During the course of this studv, material in Ixodes putus: Kishida, 19.30a: 2. the 406th Medical Laboratory Collection (No. 148) was again sent to Mr. G. M. Kohls, who Discussion: gave the following opinion: Keegan and Toshioka (1957) listed /. uriae

Very similar to /. brunneus Koch; see in the section, Questionable Distribution Rec- Cooley and Kohls 1945, p. 205. Differs ords. As noted by them, I. fuitus. which has from brunneus in characters of the scutum, been synonymizcd by Cooley and Kohls (1945) the punctations being more numerous in under uriae. is recorded by Kishida (1930a) as 162 Hni<;nAM VouNC University Science Bulletin

/«^ Jy£fc«.<-ai»_

I'"k:. 112. Ixodes turdus; feinak-. Biological Series, Vol. 15, No. 1 Ticks 163

Tn. 7nu^i-Ji4^

Fic. ll'J. Ixoilen turdu.s. nymph. 164 Bricham Voung University Science Bulletin

Ruuchi

Vic 111. /.vfi(/cv tunliis. larva. .

Biological Series, Vol. 15, No. 1 Ticks 165 occurring on a bat, Ntjctahis sp., in Niigata and Biology: Nagano prefectures. Kishida's report of putus This species is said to be a three-host tick. appears to be highly doubtful in view of the Males do not feed and are assumed to copulate fact that I. iirUic is primarily a parasite of marine with the females after they have dropped off birds. Keegan and Toshioka ) ( 1957 expressed from the host. Zumpt (1952) considers that the doubt about his identification. wide distribution for this species can be at- Neumann (1902) created the genus Cera- tributed to the sticky eggs, which have a long tixodes for putus on the basis of the structure of incubation period and can be easily transported tlie capitulum and the peculiar character of from one place to another on the feet of birds. Haller's organ. Since then, the genus Ceratixodes Other biological infonnation can be obtained has been either suppressed or used, depending from Nuttalfand Warburton (1911). on the workers. Zumpt ( 1952), having discussed the status of the genus, considered that the Disease Relationship: characteristics of iiriac were sufficient to sepa- Unknown. rate it from Ixodes. However, the taxonomic problem related to Ceratixodes does not appear Ixodes vespertilionis C. L. Koch to be settled; moreover, Japanese specimens (Fig. 115-118) were not available for examination. Therefore, Ixodes vespertilionis C. L. Koch, 1844:217-2.30, the species is here treated as belonging to the Fig. 9; Arthur, 1956:180-184, Fig. 1-13; Hoog- genus Ixodes, .sen.m lata. straal, 1956:567-573, PL 67;' Keegan and Illustrations of this species were included in Toshioka, 1957:25, PI. 39-40; Asanuma, papers by Nuttall and Warburton (1911) and 1965a: 113-116, 1965b:399, Fig. 221. Pomerantzev (1950). The following information was taken from the literature. Discussion: Arthur (1956) and Hoogstraal (1956) both Dl\gxosis: refer to a single collection (RML No. .32112) of This "puffin-tick" has very unique character- /. vespertilionis from RhinoIoj)hus ferrum- istics. The size is very large in both sexes. The eqtiinum Sawada, "Myiagi," Honshu, 13 May male has five distinct tufts of long hairs at the 1952 (record was given in personal correspon- posterior margin of the body; the 3rd palpal seg- dence from Kohls to Hoogstraal). The error in ment is conicallv shaped. In the female the body spelling (should be "Miyagi") is perpetuated is covered thicklv with white hairs; the 3rd pal- in Arthur (1956) and Hoogstraal (1956). Kee- pal segment is much broader and shorter than gan and Toshioka (1957) examined this single the 2nd, and the suture between them is poorly female specimen and gave several records of defined, the 1st palpal segment is dorsally vespertilionis from bats collected on Honshu prominent. and Shikoku. Specimens from Hokkaido were borrowed from Mr. K. Hattori, and other study Distribution and Hosts: material was collected in Iwate Prefecture and in Korea 406th Medical Laboratory personnel. The tick has a wide geographical distribu- bv tion. The recorded localities for this species comprise many islands or seashores facing Diagnosis: oceans encircling both poles. The species is This bat tick is very distinctive and may be therefore called a "bipolarlv distributed" tick. easily distinguished from other Ixodes spp. The

Zumpt ( 1952 ) discussed the distribution, cit- legs are thin, much longer than the body, giving ing the opinions expressed by Schulze ( 19.38 ) a "spidery" effect; all coxae lack spurs; the anal Detailed localities for the tick are given bv groove is round in front of the anus and the pos- Nuttall and Warburton (1911), Schulze (19.38), terior arms are nearly straight or slightly con- f:oolcy and Kohls (1945), and Zumpt (1952). verging. In the male, the palps are clavate and The hosts are mainlv sea birds inhabiting have numerous long setae; the hypostome is the areas mentioned above. They are gulls, al- pointed at the tip with faint indications of small, batrosses, petrels, shearwaters, and penguins. scalelike teeth. In the female, the scutum is nar- Pomerantzev (1950) and Nuttall and Warbur- row and much longer than wide, with the widest (in Nuttall and ton ( 1911 ) state that it occasionally attacks man. point just behind the middle The sole reliable record in the Japanese litera- Warburton, 1911, broadest in the middle; in ture is Asanuma's report (1961, 1965a) which Pomerantzev, 1950, widest in front of the mid- contains the record of a nymph and a larva from dle), the basis capituli is dorsally triangular and a puffin. Ltinda cirrhata. on Hokkaido. has a flange externally directed at the postero- Bbicham Vounc University Science Bulletin 166

jy/f///y.l yx^

lemale. Fig. 1 15. Ixodes tcspcriilioni^, Biological Series, Vol. 15, No. 1 Ticks 167

Fig. 1 16. Ixodes vexpertilionui, male. Bricham Young Univehsity Science Bulletin 168

1.0 mm

Fk;. 117. Ixdilcs vc\i)crtilioru.s. nymph. Biological Series, Vol. 15, No. 1 TiCK.S 169

2 mm ^.77U*

Fir.. 118. Ixodes vesjierlilionis, larva 170 Hricham Young University Science Bulletin external juncture; the liypostomc is lanceolate Arthur's (1956a) comparison of the data and has long denticles. from Switzerland and from Macedonia leads

him to believe that, because there is a high DiSTIUBUTION AND HOSTS: reasonable catch of partiallv and full en- The distribution and hosts of this species in gorged ticks between October and January areas other tlian Japan are presented by Arthur and a number of unfed nymphs and females

(1956) and Iloogstraal (1956). This species oc- during the summer, feeding is accomplished curs on a variety of bat hosts in Europe, Africa, mainly during the winter months. This pic- the Near East, and Japan. Russian workers re- ture, possibly modified bv the host's sea- port the tick from Russia, northern Iran, and sonal breeding cvclc and activity, re(|uires western Siberia. Hosts from which Ixodes ves- further observation. Nuttall and Warburton jwrtilionis has been taken in japan are given in (I91I) postulated that males may either Appendix 2. feed very rapidly and then leave the host, or that they may not feed at all. Neumann (I9I6) believed that the various degrees of engorgement in which male specimens are found might not necessarily prove that males do feed but rather mav be an indication of degree of nvmphal feeding. This conclusion is based on the atrophy of the male hypostome in comparison with its robust development in females and in immature stages (Hoogstraal, 1956).

Disease Relationship: Unknown.

Genus Rhiplcephalus C. L. Koch. 1844

Usuallv ornate. Palpi short. BasLs capituli usually hexagonal dorsallv. Eyes and festoons present. The males with a pair of adanal and accessory shields. Coxa T bifid. Spiracular plate comma-shaped.

Rhipiccphahts sanguineus Latreille, 1806 (Fig. 119-122)

Ixodes sanguineus Latreille, 1806:157

Rliipiccplialus sanguinctts: Koch. 1844:2.3.S-2.39; Sugimoto, 1937b:610-612; Keegan and To- shioka, 1957:26-27, pi. 42.

Discussion:

Map .36. Known distriliiitid )f Ixodes spvrtiUou Although this tick, known as the "kennel tick," "brown dog tick," or "tropical brown dog Biology: tick," is one of the most widely distributed ixo- Nothing is known of the biology of the Japa- did ticks, present infonnation indicates that it nese population. Excellent reviews of biology, is very rare, at least in Japan, Korea, and the morphology, etc., are given by Arthur (1956) Ryukyu Islands. Sugimoto ( 1937b) reported that and Iloogstraal (1956): "No males have been R. sanguineus had been taken in Kumamoto

found on bats, but thev wander over the walls Prefecture, Kvushu, but it has not been found of the caves, partieularlv where the rocks arc b\- other collectors in that area. creviced. Unfertilized females and unfed im- During the course of this stud\-, threi' lots mature forms occupy similar niches. It seems were examined and identified as R- sanguineus. likely that the fully fed stages secrete them- judging from the 406th Medical Laboratory col-

selves between the stones of the caves to digest lec-tion n>cords, it is difficult to detennine blood meals" (Arthur 1956). whether this species has been established in Biological Series, Vol. 15, No. 1 Ticks 171

Fig. 119. Rhipucpluihi.s sanguineus, female. Brigham Young Univebsitv Science Bulletin 172

Sonobe

iMG. 120. Rhipiiciihcilus sunj^uiiinis, male Bi()LO(;icAL Sehies, Vol. 15, No. 1 Ticks 173

TAmIo

Fig. 121. RhipirrphnJtis sdUf^tiiurtis. iivmpli. Bhigham Young University Science Bulletin 174

-A^

Fig. 122. Rltipiccphalus- suni^uinvus, lana. :

Biological Series, Vol. 15, \o. 1 TicK.s 175

' 1 ' •- ,- . Japan. However, it is readily siippcsed that it . ,

Rhipicephalus sanguineus , inav have been introduced into Japan on pets of ^

/ U.S. Forces personnel or in household effects.

Feldniann-Muhsani ( 1952) has suggested that the tick commonlv known as R. sangttineus is actually not one species, but two. She pro- posed the specific name secundus for the new species, which is distinguished from sanguineus s. str. by the nature of the female genital aperture, and in immature stages by the shape of the capitulum. This distinction has not been fullv accepted bv Hoogstraal (1956).

DlAGNO-SIS

Since there is only one species of the genus

Rhipiccpltalus that occurs in this area, it is easil\- identified.

Distribution and Hosts:

Leeson ( 1951 ) did not include Japan, Korea, and the R^ukvu Islands as part of the distributional area of this species, but the special map published by the American Geographic Societ}'

( 1954 ) shows Korea and the Rvukyu Islands as areas where sanguineus occurs. Anastos (1950) gave its distribution in the East Indies, and Kohls (1950) discussed its occurrence in the Philippines. As reported by Anastos (19.50) and Hoogstraal (19.56), sanguineus is primarily a parasite of dogs but also attacks a great variety of domestic and wild animals. It has been taken from l)irds as well as mammals, and in some areas has been known to attack man.

Biology:

Hoogstraal ( 1956 ) has reviewed the literature on the biology and medical importance of this tick. According to Hoogstraal's review, it is said to be a three-host tick, and the life cycle mav be completed in as little as 6.3 days under favorable conditions. If conditions are unfavorable, it may be prolonged for many months. "In Nuttall's laboratorv experiments, larvae fed on dogs and rabbits, nvniphs on dogs, jackals, and hedgehogs, and adults on dogs and jackals" (Hoogstraal, 1956). He also cited several reports on parasites and predators of this tick. Hunter-

elhis hookcri Howard, 1907 ( =IxocUphagus cau- curtci du Buvsson, 1912), a wasp, has been oc- casionallv reported as a parasite of R. sanguineus in .\frica, Brazil, USA, and other areas of the world. The eggs of this wasp are laid in the body of the nvmph and the developing hymenopter- ous larvae feed on the contents of the engorged nvmph and pupate in the body. A predaceous spider, Teutena triangulosa Wick, has been observed feeding on both immature and adult forms )

176 BiiK.iiAM VouNC Univebsity Science Bulletin

Cijclemijs flavomarginatd by N. Nakayama. He (1907). Although Pomerantzev (1950) stated did not indicate the date and locality, but the that the species had been recorded from Japan, host turtle is found on Ishigaki (Is.) and Irio- such records do not (K-cur in the Japanese litera- mote (Is.) in the Hyukyus. ture. He pn)b;ibl\- cited Neumaim's record. Nutt- Ambhjonima <^eoei)it/(lae (Cantor, 1847) is all and \\';irl)urton (1915) expressed doubt as recorded from Ctjcleimjs fkwoimirginata on Irio- to the correct identification of Neumann's ( 1907 mote (Is.). specimen from a horse in Aomori Prefecture, Japan. There are no specimens in Japanese col- Ambh/oynina ciiprhim Neumann, 1(S99: 2 19-221;

lections at the present time, and the species has i Nakatsuji, 19-13: lcS2-lS4, Fig. 1. not been collected by Japanese acarologists. Nakatsuji (1943) reported the collection of one female tick of this species from a turtle, Ilaeniaphysalis shibutamiensis Yajima, 1955. Geoeimjda spengleri, on Okinawa. The figures This is apparently a nomen nudum. Although published with this report are not adecjuate for Keegan and Toshioka (1957) state that Yajima discrimination. described the species in the ]aj)anesc Jattrnal

Anastos ( 1950 ) stated that there were no au- of Sanitary 7.oolofiy 6( 1 ) :52-53, the specific thentic records of parasitism of reptiles by A. name was neither given in the paper nor referred ciiprhim cijprium; it may be that the specimen to in later papers bv Yajima. Tvpe specimens reported bv Nakatsuji was actually '^eoemijdae, were not available for examination. v\'hich is known to occur on turtles in the Ryu- Haemapliysalis u(itamil>ci Yajima, 1942. (See kyu Islands (Keegan and Toshioka, 1957).

' under //. flava. ) Haemaphiisttlis cinmiharina C. L. Koch, 1S44: llaemaplii/salis hirudo L. Koch, 1877. 337; Kishida, 1922a ;846-855. There were no figures with the original des- Kishida (1922a) listed both sexes of this cription, and the description itself was not ade- species in his keys to the species which occur in (juate for detennination. See under //. concinna. Japan. Keegan and Toshioka ( 1957 ) stated that the occurrence of this species in Japan is very Hyalomnm ac<^ii])ticum (Linnaeus, 1785); Kish- unlikely. Yajima (1956) listed it a,s cK-curring ida, 19.36:139. in Aomori Prefecture, but his record may have Kishida (1936) reported this species from a been derived from Kishida ( 1922a ( see Ap- ) cow on Quelpart Island ( = Chejudo Island, pendix Keegan and Toshioka (1957) also 1). Korea), and it is the onlv record of this species ({noted Kohls"s 1957 personal communication area. indicates that from this Hoogstraal (1956) j noting that cinmiharina, described from Brazil, area //. aegypticum occurs in the Mediterranean | is for chordeilis, in a valid name which occurs and the Near East. It has been recorded from North America. Russia, many parts of Asia Minor, and Afghan- Haema]>htjsalis jxipuana Thorell, 1882:62-66; istan. This species is primarily a p;uasite of tor-

attach other reptiles as well , Nakamura and Y;ijima, 1937:158-159, Fig. toises but may to as birds. Pomerantzev ( 1950) i 5-9 in PI. VIII. mammals and gave infonnation concerning the distribution and Nakamura and Yajima ( 1937 ) reported two biology of this species. On the basis of this in- female specimens collected from a dog at Hamg- is doubt concerning the fonnation, there some ] yong-pukto. North Korea, 26 Julv 1935. Accord- validity of the record from Quelpart Island (Kee- ing to Anastos (1950) this is essentially a tropi- gan and Toshioka, 1957). cal species, and Kohls (1957b) expressed doubt that this species occurs in Korea and other non- \xodcs acumimilus \vumd\m, 1901:287; Ki.shida, j tropical regions. The illustrations provided by 19.%: 142. Nakamura and Yajima (1937) do not look like "Kishida ( 1936) stated th;it Mr. Ak:isawa had , those given by Anastos (1950) but appear to be collected Ixodes acuminutus from central Korea similar to those of //. japonica. in 1928. No specimens have been reported from Haemaplnjsalis punctata Canestriui and Fanzago, Korea, Japan, or the Ryukyu Islands since that 1878:189. date, and the species is not represented in exist- ing collections from these areas" (Keegan and llaemaplujsaliH cinnahaiiud jiuiuialir. Neumann, To.shioka, 1957). 1907:215-2.32, Fig. 1-4; Kishida, !922a:852; Pomerantzev, 1950:92-93. Ixodes affiuis Neuiiumn, 1899:120; Kishida,

Kishida (1922;i) listed l[. c. punctata in his 1930a:3; Suzuki, 1930:90-92. Fig. 1.

/. from ;i rabbit, key. which seems to be derived from Neumann Kishid;i ( 1930a) listed affinis Biological Series, \'oi.. 15. \o. 1 Ticks 177

Lepus hmclii/urus an^tistidens. at Fukushinia it later as a new species (1930a). According to 15 City. Deccniher 1929. collected bv Dr. Ohara. Keegan and Toshioka ( 19.57), the type specimens Suzuki (1930) described and illu.strated the have been lost and no other material was avail- species using a male specimen which had been able for study. Judging from the host and the taken from a rabbit of the same species. These figures given by Kishida (1922b), the species specimens were not a\ailable for study, and appears to be similar to Ixodes persulcatus. unfortunatc]\' the reproduction of Suzuki's illus- Ixodes putus (Pickard-Cambridge, 1878). See tration of the male is not satisfactory for deter- under /. uriae. mination of the species.

Ixodes corcensis Kishida, 193.3; Kishida, 1936:139. Xenoacarus Kishida, 1925. "Efforts to locate the original description of Palpi short; scutum not heavily sclerotized; this species were not successful. Kishida (1936) postanal median groove extends to posterior body referred to this tick, giving the tvpe locality as margin; festoons and eyes absent. suburbs of Keijo = Seoul), the host as ( and Xenoacarus ut'ikinoi Kishida, 1925:31. Nyctereutes koreensis. The species is not repre- The original description was not avail- sented in any of the collections in Japan, and Dr. able for translation. Dr. Kishida infonned Kishida has infonned the author that the holo- Dr. Seiichi Toshioka that the original des- type was lost during the war" ( Keegan and To- cription consisted of only a few lines and no s'hioka, 1957). illustrations. No specimens were available Ixodes fossulattis Neumann, 1899:120; Kishida, for study. The generic diagnosis given above 1930a: 2. and the following species description were

Kishida ( 19.30a ) listed this species as occur- taken from a discussion of the species in ring on cattle and dogs at Sefuri-muri. Kanzaki- Essential Para.ntologii of the Domesticated gun. Saga Prefecture. Keegan and Toshioka Animals. April 1926," p. 199-200, by S. Yo- (1957) state that the specimens are not in the shida and T. Makino. A figure with this collection of Dr. Kishida and no additional discussion showed a dorsal view of the specimens have been collected by Japanese scutum and capitulum of the female of workers, /. fossulattis is originally an Ecuadorian makinoi. species. "Male: Unknown.

Ixodes Juiiij)imis Yajima, 19.55; Keegan and To- "Female: Postanal median groove ex- shioka, 1957:31. tends to posterior body margin; dorsal sur- basis capituli rectangular and bow- This is a nomcn nudum. The specific name, 7. face of anteriorly. Dorsal scutimi not hapjiinus. is not given in any published papers ed forward heavily sclerotized and monochromatic. Pal- by Yajima. See under /. acutitarsus. pi short as in Haemaplnjsalis. Coxa I with a Ixodes haijashii Nakatsuji, 1942:287-328. Fig. 3. large spur. [One unintelligible phrase pre- a "This species description was based upon ceded this statement.] Festoons and eyes single female tick taken from a dog on Miyake absent. Legs and scutum yellowish in color, Island. 26 July 19.37, by K. Hayashi. Figures even in engorged specimens." Tliis species given of the female include dorsal and ventral has been found on leaves or stalks of plants. the posterior aspects of the tick (showing only It has been collected from horses, etc. Para- portion of the capitulum), hypostome, and tar- type collected from Japan: Honshu, Aomori sus I. Unfortunately these are not adequate for Prefecture, Kamikita-gun, Misawa-mura, identification of the species. The hypostome, as Hirosawa pasture, August, 192-5 (Keegan figured, is unlike that of any of the Japanese and Toshioka, 1957). Ixodes. No specimens of this tick were collected by 406th MGL personnel and the species is not represented in an\- of the Japanese collections" ACKNOWLEDGMENTS (Keegan and Toshioka, 1957).

Ixodes lisphilus Kishida, 19.30; 1922b: 962-964, We are happy to express our appreciation Fig. 1-2, 19.30a:.3-4. to Mr. Glen M. Kohls, Rocky Mountain Labora- Hoogstraal, This species was described by Kishida tory, Hamilton, Montana; Dr. Harry (1922b) using nvmphal specimens taken from a U.S. Naval Medical Research Unit No. 3, Cairo, Dr. Shigeo Kitaoka, National Institute squirrel, Sciunis lis. Nagano Prefecture, Japan, Egypt; of Animal Health, Kodaira, Tokyo, who have 1 June 1918, by Uchida. However, he (1922b) identifica- did not give the specific n;une but designated been very generous in confinning our .

178 Bhigham Vounc University Science Bulletin

tions or in loanins^ valuable specimens and ofter- Okumuni, Toyonaka-shi.

iu'Z advice on taxa, the taxonoinie status ot whicli We are also grateful to the following persons. was in (juestion. Department of Kntomologv, 406th Medical Lab- We wish to express our appreciation not oratory, USAMCJJ, who have prepared illustra- only to the above but also to the following tions for this publication: persons who have cooperated in various ways Mr. Saburo Shibata, Mr. Takashi Ando, Mr. to make possibU; the completion of the paper: Shozo Ohtawa. Mr. Masao Ilasunuma, Mr. Yii- Dr. Zen-emon Ono, Hokkaido Institute of saku Sonobe, Mr. Yoshitsune S;isaki, Miss Mut- Public Health, Sapporo; Dr. Kiyoshi Asanuma, suko Misaki, and Mrs. Atsumi Ryuchi. In addi- Research Institute for Natural Resources, Tokyo; tion, we express appreciation to Miss Yoshiko Mr, Keisaku Tattori, Hokkaido Institute of Public Yoshida for her careful recording and filing of Health, Sapporo; Mr. Kimio Endo, Iwaizumi-cho, the original illustrations made by the above art- Iwate Prefecture; Mr. Kimito Uchikawa, Nagoya ists. Institute of Public Health, Nagoya, for the kind We express thanks to Mr. Hiroshi Suzuki and loan of specimens. Mr. Shogo Sasagawa for their cooperation and Dr. Hiroshi Takahashi, Medical School, Tok- devotion to dut\' in the rearing of field material yo, Japan Ground Self-Defense Force; Dr. Kazu- for laboratory life-cycle studies and to Dr. Kazuo yuki Uchinomi, Hachijo Station of Tokyo Cent- Tanaka for hLs continuous encouragement and ral Health Center; Dr. Naoki Namba, Hokkaido timely advice on taxonomic problems. National Argicultural Experiment Station, Sap- Mr. Kiyoyuki Mizusawa and other field col- poro; Dr. Ryuichi Taniguchi, Shintoku Experi- lectors of the 406th Medical Laboratory provided mental Station of Domestic Animals, Shintoku, a large proportion of the material examined in Hokkaido; Mr. Koichi Ono, Mita Pasture, Koz- this paper. awa, Hokkaido; Mr. Toshihide Takeda, principal We wish to take the opportunity to express of Yamato Middle School, Aniaini Oshima, Ka- thanks to LTC Alexander A. Hubert, chief of the goshima Prefecture; Mr. Zentoki Hajime, Yuwan, Department of Entomology, and Dr. George M. Amami Oshima, Kagoshima Prefecture; Dr. Shoi- Davis, Department of Medical Zoology, for their chiro Ohara, Ohara Hospital, Fukushima; Mr. frequent encouragement and for reviewing the Susumu Yamamoto, Kagoshima Institute of Pub- manuscript; to Mrs. Anne Lester for her pains- lic Health, Kagoshima; Dr. Kunitada Sato, Shiz- taking tvping and correcting of early drafts of unai Pasture of Hokkaido University, Shizunai, this manuscript; we acknowledge the continuous Hokkaido; Dr. Yasusuke Nagayoshi, Fukushima support and interest of COL J.E. McCarty, Com- Hospital, Kushima, Miyazaki Prefecture; Dr. Me- manding Officer of the 406th Medical Labora- guinu Hasegawa, Hokkaido Institute of Public tory. Health, Sapporo; LTC Yasuo Nakamiya, Camp We express our thanks to Mr. Robert C. Obihiro, Japan Self-Defense Force, Obihiro Hok- Saunders and Mr. Lynden P. Baum for their kaido; Dr. Hajime Kanio, Department of Parasit- review of the manuscript and Miss Phyllis Mad- ology, Tottori University, School of Medicine; den and Miss Dianna Rice for typing the final Dr. Atsuo Sato, Department of Parasitology, Ka- draft of the manuscript. All are students at goshima University; Dr. Asahiko Yajima, Kitama- Brigham Young University. chi. Kichijoji, Musashinoshi, Tokvo; Dr. Yutaka The authors would like to acknowledge with Saito, Department of Medical Zoology, Niigata sincere appreciation all others who may have University of Medicine, Niigata; B. M. Mutsuko been omitted by name but who gave assistance.

LITERATURE CITED

American Geocrapiiicai. Society. 1954. .Atlas of of lick from Austr.ilia. J Parasilol. 41 ( 1 ) : 18-23. di.seases. Plate 12. World distrihiition of rickettsial Arthur, 1). R. 1956. The Ixoilcx ticks of Chiroptera

- Parasit. : 180-196. di.sea.ses tick and mite vectors. ( New ^'ork ) (Ixodoidea, Ixodidae). J. 42(2) Amerson, a. B.. Jr. U)68. Tick distrihulioii in the .Ahihuh. D. R, 19,57. Studies on exotic ticks (Ixodoi- Central Pacific as influenced bv sea bird movement. dea. Ixodidae) from United States Navv and Army :332-.3.39. :681-694. J. Med. Ent. 5(.3) activities. |. Parasit. 43(6) Ana.stos, G. 1950. The scutate ticks, or Ixodidae, of .\inTiuR, D. R. 1960. Ticks, a monograpli ol the

Indonesia. Kntomologica Americana 30( 1-4 ); 1-144. Ixodoidea. Part V. On the genera DcrmiucntOT, Anonymous. 1964. Survey on ticks in the pastiire at Anoretitor, Coxmiommu. Bonphiliif:. and .\/(;rp;r();)i(.s. Sanbe. Sliimane Prefecture. Report of the Institute C;ambr. Univ. Press. 251 p. of Animal Health of Sliimane Prefecture, p. 1-10. Ahthuh, D. R. 1963. British ti

Biological Series, Vol. 15, No. 1 Ticks 179

Mag. 53(6):306-307. (Japanese te.\t.) macy), p. 181. (Japanese text.) As.ANUMA, K. 1942. On tlie dog tick Hucmupht/salis .\sANUMA, K. 196.5a. Suborder Ixodides, p. 101-128.

campunuliitii. Ciinvciken 11 ( 1 ): 17-26. ( |apanese In Sasa, Mites, an introduction to cliissification, text. bionomics and control of Acarina. Univ. of Tokyo .As.\NUMA, K. 194-4a. Some obsenations on the (luan- Press. 486 p. (Japanese text.)

tity of blood sucked by i.xodids and on their body AsANUMA, K. 1965b. p. 397-401. In Okada et al., weight. ( Studies of the egg-laying capacity of New illustrated encyclopedia of the fauna of Japan, Ixodidae. I.) Inst. Re.s. Sci. Re.sour."Proc. (6):35- Vol. II. Hokuryukan Co., Tokyo. 803 p. (Japanese 40. (Japanese text.) te.xt. As.\NUMA, K. 1944b. Minimum quantity of sucked AsANUMA, K. 1969. Personal communication. blood capable of inducing egg-deposition in ixodid-s. AsANUMA, K., AND S. FuKUUA. 1957. On the life-his-

( Studies on tlie egg-laving capacity of Ixodidae. of a tick. Ixodes signatus, infesting a black-tailed 11.) Inst. Res. Sci. Resour. Proc. (6):41-46. gull (Larus crassirostris) in Sanit. (Jap- Japan. Jap. J. iuiese text. Zool. 8(3):147-1.59. (Japanese text. AsANUMA, K. 1944c. Influence of the (juantity of AsANUMA, K.. AND K. KosAKA. 19.54. Notes on the oc- sucked blood on the number of eggs laid by ixodids. currence in Japan of a tick, Hacmaphi/satis welling-

( Studies on the egg-laving capacity of Ixotlidae. toni, collected from a Japanese gray thrush, Turdus III.) Inst. Res. Sci. Resour. Proc. (6):47-54. (Ja- cardis cardis. Misc. Rep. Res. Inst. Nat. Resour. panese text. 36:104-107. (Japanese text English translation: .\sANUMA, K. 1944d. Egg-laying habits of ticks. NAMRU 3-T35.) Misc. Rep. Res. Inst. Nat. Resour. (19):21-31. AsANUMA, K., AND K. KosAKA. 1955. Notes on a tick, AsANUMA, K. 1947a. Note on the role of blood in Ixodes turdus Nakatsuji 1942, found on birds in ovulation in a tick, Hacmaphysalis hispinosa, to- Japan. Bull. Biogeogr. Soc. Jap. Appendix 16-19: gether with remarks on the so-called specific dif- 192-196. (Japanese te.xt. English translation: NAM ferences in the number of eggs laid by ticks. RU 3-T50.) Studies on the ovipositing ability of ticks, V. AsANUMA, K., AND H. Nakacawa. 1955. Studies on Seibutsu 2(1):7-14. (Japanese text.) endemic and medically important arthropods on AsANUMA, K. 1947b. p. 971-974. In Uchida et. al.. Niijima Island, Tokyo-to. Ochanomizu Igaku Zasshi

Rev. ed. of illustrated encyclopedia of the fauna 3( 6 ) :549. Japanese text. EngUsh summary, 680. ( p. of Co., Asanuma, K., K. Okubo., and S. Fukuda. 1955. In- Japan. Hokur^ukan Tokyo. 1898 p. ( Jap-

iinese text. festation of a black-tiiiled gull ( Larus crassirostris .\sANUMA, K. 19.51. Notes on the tick, Ixodes angus- with a tick, Ixodes signatus, in Kabushima, Aomori tus Neuman, new to Asia. Misc. Rep. Res. Inst. Prefecture, Japan, Misc. Rep. Res. Inst. Nat. Resour. Nat. Resour. 22.1-4. (Japanese te.xt. 38:85-86. (Japanese text.) AsANUMA, K. 1954. Studies on ticks and bloodsuck- Asanuma, K., and N. Sakural 1958. On the seasonal ing mites infesting rats (genus Rattus) in Japan. occurrence of the tick, Haemuphijsalvi flava, on

Part I. Misc. Rep. Res. Inst. Nat Resour. ( 36); 108- wild hares (Lepus timidus bracht/urus) in the en- 119. (Japanese text. English translation: NAMRU demic area of yato-byo or tularemia in Chiba Pre-

.3-T48. fecture, Japan ( A preliminary report ) . Misc. Rep. AsANUNLA, K. 1955;l On the i.xodicLs and laeapitids Res. In,st. Nat. Resour. (48):28-39. (Japanese text. pariisitizing mice of Hokkaido, and mode of trans- English summarv, English translation: NAMRU mission of pathogens among liost mice bv the acari. 3-T46. Nihon I ji Shinpo (1611):! 240- 1242.' (J apanese Asanuma, K., and N. Sakurai. 1960. Comparison text.) between the infestation of Ixodes japonensis and .\SANUMA, K. 1955b. Mode of disseminating the caus- Haemaphi/salis fliiva on wild rabbits. 7. Rep. Ohara ati\'e agent ;imong the rats with various ticks and Hosp. 4(2):26. (Japanese text.)

) :1240-1242. AsANUM.'\, K., N. S.\kuhai, S. Ohara, H. Odajima, K. mites ;ls vectors. Jap. Med. J. ( 161 1 (Japanese text.) KosAKA, and H. Nakagawa. 1955. On the host AsANLMA, K. 19.56a. Tularemia, ticks, and mesostig- and distribution records of a tick, Haemaphysatis matid mites of the Far Eiist. Ann. Prof. Rep., 406th flava, a dominant panisite of wild hares in Japan. Medical General Laborator)'. p. 91-96. Mi.sc. Ref. Res. Inst. Nat. Resour. (.37 ); 127-128. .\s.\NUMA, K. 1956b. Personal communication. (Japanese te.xt. English translation; NAMRU 3- AsANUMA, K. 1957. Occurrence of the tick, Ixodes T49.) angustus Neuman. in Hokkaido, Japan, with re- .\sanuma, K. and K. Sekikawa. 1952. Studies on

marks on its larval and n\mphal stages. Jap. |. ticks of the genus Ixodes parasitic on small rodents Saint. Zool. 8(4):2.34. (Japanese text.) of the superfamily Muroidea in tlie Japanese Islands .\sA>aiMA, K. 1958a. New hosts of ixodid ticks. Shawa (a preliminarv note). I. Misc. Rep. Res. Inst. Nat.

.3.3rd Monbusho Sogo Kenkvii Hokoku Shuroko Resour. ( 28)';107-1 16. (Japanese text.) (Med. and Pharmacy), p. 279. (Japanese text.) Asanuma, K., .'Vnd K. Sekikawa. 1953. Studies on AsANUMA, K. 19.58b Ecology of Ixochdae parasitic ticks of the genus Ixodes parasitic on small rodents on media animals for yato-b\o. 2. Rep. Ohara Hosp. of the superfamilv Muroidea in the Japanese Islands 2(1):23. (Japanese te.xt. (a prehminary note). II. Misc. Rep. Res. Inst. Nat. .\sANUMA, K. 1958c. Yato-bvo in Sado Island. 2. Resour. (31):99-112. (Japanese text.) 1960. Rep. Ohara Hosp. 2( 1 ) :.38'-39, (Japanese text) Asanuma, K., A. Shishido and M. Kitaoka. .\sANV>L'^, K. 1960. Host and distribvition records of The hfe of Rickettsia in the lx)dies of larvae and the soft tick, Ornithodoros capensis Neumann, in nymphs of IIaenuiphysatis- tiispiiwsa. 7. Rep. Ohara Sanit. text.) Hosp. 4(2):26. (Japiuicse text.) Japan. Jap. J. Zool. 11(2):94. (Japanese .\sANUMA, K. 1961. Blood-sucking acari - 2. Para- .\sANUMA, K., T. Takeda, and H. Nakacawa. 1961. birds in the sitic ticlts of sea birds in Japan. Showa 36tti Mon- Some observations on ticks found on Fuji area. (Japanese abstract of report before busho Kenkyu Hokoku Shuroku ( Med. and Phar- Mt. )) ) . ) )) ) ) )

ISO Bmr.HAM VouNc; University Science Bulletin

Anastos. 1966b. Ixodid ticks 13 Ann. Meet. Jap. Sw. Saiiil. Zool., |ulv 5.) Elbl, A., and G. Vol. II. Sanit. Zool. 12(2): 139. (Acarina, Ixodidae) of central Africa. Jap. J. 1795. Ann. Mus. R. Afr. Asanuma, Y. 1943. Con.siderations on the amount of Cienus Ixodes Latreille. s. Sci. Zool. (146). 205 p. 1>1o

Cooley, R.' a. 1946. Note on the tick Ixodes Hoocstraal. H. 19,56. .\frican Ixodoidea. I Ticks of angustiis Neumann. J. Parasit. 32(2):210. the Sudan (with special reference to Ef|uatoria CooLEY, R. A., and G. M. Kohls, 1945. The genus Province and with preliminary reviews of the Ixodes in North America. Nat. Inst. Hlth Bull. genera Boophilus, Margarojtus, and Ht/idonima). 184:1-246. Bureau of Med. and Surgers . Department of the Dhanda, v., and U. R. Biiai. 1968. Ilacinuphi/sidi.s- Navy. 1101 p. (Haemap]ujsulis)indoflmii sp. n. (Acarina: Ixodi- HooosTR^Ai.. H. 1961. Rediscriplion of Ilnermiplii/-

dae) from the Himala\an region of Uttar Pradesh s(dis ( Alloccriiru) incrmis avouommoides War-

and Madras. India. Parasit. .54(5) : 1063-1067. Parasit. J. burton. 1913 (Ixodoidea, Ixodidae). J. 47 Donit-z, \V. 1905. Die Zecken des Rindes als Kran- (2):317-318. kheitsubertrager. S. B. Ges. Naturf Fr. Berlin H<)Oc.sTR\Ai., H. 1962. Haemaphijsahs ncpidensis sp. 4:105-134. n. from a Himakuan rodent and man and de- Elbl, A., and G. Anastos. 1966a. Ixodid ticks scription of the male of H. aponommo'des War-

(Acarina, Ixodidae) of central I. Africa. Volume burton (n. comb.) (Ixodoidea Ixodidae). J. Para- General introduction. CJenus Andili/omma Koch, sit. 48(2): 19.5-203. 1844. Ann. Mus. R. Afr. Cent., Ser. .Svo. s. .Sci. Hoocstraal, H. 1966. Ticks in relation to human Z

BiOLOOiCAL Sehies, Vol. 15, No. 1 Ticks 181

3()S. Karsgh, F. 1880. Vier neue Lxodiden des Berliner HooGSTK.\AL, H. 1967a. Tick - Iwme hemorrliagic Museums. Mittheil. d. Munchener Entomol. Ver- fevers, encephiUitis, and typhus in USSR and eine. Jahrg. 4:141-142. southern Asia. Exper. Parasit. 21( 1 ) :98-lll. Katsura, S. 1957-1959. Studies on the bacterial Ht)OGSTR.\-\L, H. 1967b. Ticks in rehition to liuman and infectious diseases transmitted by ticks in diseases caused bv rickettsia species. Ann. Rev. Japan. Showa 32iul, 33rd, and 34th Monbusho Ent. 12:377-420. Sogo Kenkyu Hokoku Shuroku. p. 265-274 (1957); Hi)(k;sth.\al, H. 1968. Personal communication. p. 277-288 (1958); p. 327-336 (1959). (Japanese HoocsTR.\.4.L, H. 19e9a. Letter No. 1013. 28 Mav text. 1969. N.WIRU 3. Kaw.^vshima, K. 1963. Studies on the acarine fauna HoocsTRA.^L, H. 19(39h. Hacmaphiisulis (Allocenwd) of the Ryukyu Islands ( 1 ) . Some records of ticks. kitaokai sp. n. of |ap;ui, and keys to species in the Jap. J. S;uiit. Zool. 14(2): 103. (Jap;inese text.) stnicturidly primiti\e subgenu* Alloceraea Schulze K.\WASH1MA, K., H. KaMO, and I. MiYAZAKI. 1960. A

of Eurasia ( l.xodoidea, I.xodidae). J. Parasit, 55(1): case of human infestation with a hard tick, Am- 211-221. bhjomma tcstudinarium in Koch, Japan. Kyushu J. HoocsTKAAL, H., AND M. N. Kaisek. 1960. Booplii- Med. Sci. ll(2):77-80. liis kohlsi n. sp. (,'Vcarina: Ixodidae) from slieep Keegan, H. L., and S. Toshioka. 1957. Ixodid ticks

goats in |ord;ui. Parasit. ) and J. 46(4 :441-448. of Japan, Korea, and the Rvukyu Islands. Report HoocsTK.\.\L. H., F. H. S. Roberts, G. M. Kohls, and —406th Medical General Laboratory. 37 p., 42 pi. V. Tipton. U)68. Review of Hciemaplii/sali.i Kierans, E., J. J. and G. M. Kohls. 1970. Ixodes

( Kaiserianu ) loiif:,con>h Neumann ( resurrected ) of philipi n. sp. (Acarina: Ixodidae), a sea bird Australia, Zealand, Caledonia, Fiji, in New New parasite Japiui. J. Med. Ent. 7(6) :725-726. Japan, Korea, northeastern China, and USSR, and KisHiDA, K. 1922a. Notes on the ticks of the its parthenogenetic and bisexual pop ulations famiK' Ixodidae parasitic on Japanese mammals

(Ixodoidea, I.xodidae). Parasit. : J. 54(6) 1197-1213. (Artiixlactvla: Bunodontia ) . Zool. Mag. 34(408): HocxJSTHAAL, H., AND H. Trapido. 1966. Redescrip- 846-857. (Japanese text.)

tion of the type materials of Hacmaplnjsalis ( Kai- KisHiDA, K. 1922b. Tick from a squirrel. Zool. Mag. seriana) bispinosa Neumann (India), H. (K. 34(410):962-964. (Japanese text.) neumanni Donitz (Japan). H. (K.) higrangei Lar- KisHiDA, K. 1925. Abstracts of the Seventh meeting rousse (Vietniun), and H. (K.) i/eni Toumanoff of the Tokvo Parasitological Society, No. 2459:31.

(Vietnam) l.xdidae). lit (Ixodoidea, J. P;\rasit. .52(6): KisHnJA, K. 1927. Acarina, p. 985-986. Oka et al.. 1188-1198. Illustrated enc\clopedia of the fauna of Japan. HoocsTRAAL, H., H. Trapido, and C. M. Kohls. 1965. Fig. 1897-1899. Hokuryukan Co., Tokyo. (Japan- Studies on soutlieast Asian Hacmaphifsalis ticks ese text. (Ixodoidea, Ixodidae). The indentity, distribution, KisHiDA, K. 1929. Supplements to the knowledge and hosts of //. (Kaiscriami) lu/stricis cattle tick, Supino. J. of Japanese Boophilus annulatus cau- Par:isit. 51(3):467-480. datus. Lansania 1(2): 19-22. Japiuie.se text. ( HoocsTRAAL, H., AND N. Vamac.uti. 1970. llacma- KisHiD.^, K. 1930a. Notes on Japanese ticks of the

physalis ( H. ) pentalagi Pospelova-Shtrom, a para- genus Ixodes Latreille, 1795. Lansania 2(ll):l-5. site of the (apiuiese black rabbit; rediscription Japanese text. of ( the male and description of the femiile, nymph, and KisHiDA. K. 1930b. Occurrence of a turtle tick, larva Parasit. Artd)h/omnm eordiferuin, in (Ixodoidea, Lxodidae). J. 56(2):367- Loo-choo. Lansania 374. 2(19): 134. (Japanese te.xt.) Inatomi, S., and N. Yamaguti. 1960. On an Ar- KisHiDA. K. 1935. Notes on the Acarina-mites and

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Jap. Soc. Sanit. Zool. 15:17-18, (Japanese text.) August 1935. Lansania 7(69) : 129-144. (Japanese IsHiHARA, T., AND S. IsHii. 19.56. Studies on piro- text.) pla.smosLs bv Theileria mutans in Japan. Abst. KisHiDA, K. 1936. Notes on the Acarina-mites and Proc. Mon. Meet. Staff Natl. Inst. Anim. Hlth. ticks, from the peninsula of Korea and the island 3:41. (Japanese text.) of Quelpart collected in August 1936. Lansania Itagaki, S., R. Noda, and T. Vamaguchi. 1944. 8(79): 131-144. (Japanese text.) Ticks parasitic on domestic animals of Asia. Jap. KisHiDA, K. 1939a. i3n the cattle ticks of the Horse-Racing Assn. 97 p. (Japanese text.) Boophyleae from Japan. Zool. Mag. 51(7) :.538-552. Itagaki, S., R. Noda, and T. Yamaguchi. 1959. (Japanese text.) Studies on the ticks of the domestic animals in KisHiDA, K. 1939b. Arachnida of Jehol: Order Aca- the Far East. Jap. Soc. Promotion Sci. 118 p., 27 pi. rina. Rep. 1. Sci. Exped. Manchoukuo, Sect. 5, Ito, S. 190.3. Ixodes riciniis. Chugai Iji Shinjjo Div. 1, Pt. 4, Art. 13. p. 1-49. PI. 1-.5. (561), 23(15):ia32-10,34 (p. 24-26). KtsHiDA. K. 1947. Acarina, p. 972-975. 7;i Uchida Ito, Y.. T. Komatsu, and A. Suzuki. 19.59. Chronic et al.. Rev. ed. of Illustrated encvclopedia of the tularemia and the epidemicological mass survev fauna of Japan. Fig. 2767, 2774, 2775. Hokuryii- in Takada Citv, Iwate Prefecture. 4. Rep. Ohara kan Co., Tokvo. 1868 p. (Japanese text.) Hosp. 3(l):25-29. (Japane.se text. English sum- KisHiDA, K., AND K. AsANUMA. 1965. Acarina, p. 397. mary.) In Okada et al.. New illustrated encyclopedia of Iro, T., and Y. Saito. 19.54. On the rickettsioid the fauna of Japan. Hokiirvukan Co.. Tokyo. 803 micrtxjrganlsms foimd in Steatoniissus sp. and p. (Japanese text.) Arnii.s sp. parasitic on PinistrcUus ahramus. Tokvo KiTAOKA, S, 1961a. Physiological and ecological IJLshinishi 71(10):.3-4. (Japanese text.) studies on some ticks. IV. Physiological stage and Ka.mo, H. 1962. Parasitic acari in western part of lipid deposit during the blood-sucking process in Japan. Showa .37th Monbu.sho Kenkvu Hokoku the tick. Nat. Inst. Anim. Hlth. Quart. l(2):8.5-95. Shnroku (Med. and Pharmacv). p. 173. (Japanese KiTAOKA, S. 1961b. Physiological and ecological text. studies on some ticks, V. Nitrogen and iron ) )) .

182 BmcHANr '^oUNG Univebsity Science Bulletin

excrt'tion and amount of blood meal ingested Koch, C. L. 1887. Japanesische Araclmiden und durini; tlu' hlood-siitking process in tlie tick. Nat. Myriapoden. Verh. k. k. zoolog-botan. Ges. Wien. Inst. Anim. Hltli. yuart. 1(2);96-104. 27:786. KiTAOKA, S. 1961c. Physiological and ecological Kohls, C. M. 1950. Ticks (Ixodoidea) of the studies on some ticks. V'l. Rate of digestion of Philippines. Nat. Inst. HUh. Bull. 192:1-28. l)l(K)d meal and nitrogen, iron, and sterol economy Kohls, G. M. 1957a. Acarina: Ixodoidea. In.sects during the oNipositing process in the tick. Nat. of Micronesia 3( 3) :8.5-I04. Bemice P. Bishop Inst. Anim. Hlth. ^uart. 1( 2) :1()5-1 12. Museum, Honolulu, Hawaii. KiTAOKA, S. 1961d. Physiological and ecological Koiiis, G. M. 19.57b. Ticks (Ixodoidea) of Borneo studies on some ticks. VII. Parthenogenetic and and Malaya. Malaysian Parasites — XVIII. Stud. hise.\ual races of Haetnupht/suli.s hiiintwaa in |apan Inst. Med. Hes. Malaya 28:65-94. and experimental crossing between them. Nat. KoHi.s, (;. M. 1967. Personal communication.

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fiifi.saria sp. n. ( I.xodoidea, I.xodidae), a cattle Bull. Ent. Res. 42( 1 ): 123-124. |xxrasite in Japan. Nat. Inst. Anim. Hlth. Quart. Linnaeus, C. 1758. Systema Naturae. Regnuni Ani- 10(2):25-33. mals, 10th ed. KiTAOKA, S.. AND T. MoRii. 1963. Supplementary LouNsiiURY, C. P. 1905. Habits and peculiarities of tests on the effects of new organophosphonis some South African ticks. Brit. Assn. Advanc. Sci.

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1901, with description of immature stages. ( I.xo- MiNNiNC, W. 1936. Zur Kenntnis des genus Bo- doidea. I.xodidae). Nat. Inst. .-^nim. Hlth. Quart. ophilus Curtice, II. Z. Parasitenk. 8(3) :365-370. 7(3): 145-152. Morel, P. C. 1963. Identite d'lxodes ovatus Neu- KiTAOKA, S., AND T. MoRii. 1967b. Effects of gamma mann, 1899 (Acariens, Ixodidae). Ann. Parasit. radiation and chemosterilants on the cattle tick, Hum. Com. ,38(6) :92.5-928. Boophilus microplus. Jap. I. Sanit. Zool. 18(2-3): NADCHATRAXf. M. I960. Notcs on rearing Malayan

126-129. (Japanese text.)' ixodid ticks ( Acarina. Ixodidae ) with special ref- KiTAOKA S., AND Y. Saito. 1967. Ixodes nipponetvtis erence to Ixodes firunukitus Supino and Amhli/om- n. sp. (Ixodoidea. Ixodidae). \ common cattle tick ma neoevufdac (Cantor). Malaysian Parasites — in Japan. Nat. Inst. Anim. Hlth. Quart. 7(2): .XLVIII: Stud. Inst. Med. Res. Malaya 28:217-224. 74-83. N.\CAHANA, M., AND K. Matsuo. 1962. Two species KiTAOKA. S.. AND A. Vajima. 19.58a. Ph\siological of ticks parasitic on the human body. Med. and and ecological stiulies on some ticks. I. Process Bio. 62(5):1I9-12I. (Japanese text.) of growth bv blood-sucking. Bull. N:U. Inst. .iKnim. Nakamura. T.. AND A. Yajima. 19.37. On moipho- Hlth. 34:1.35-147. logy and distribution of the family Ixodidae in KiTAOKA, S.. AND A. Yajinia. 1958b. Physiological Japan, particularly species parasitizing cattle and

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HioLocicAi, Sehies, Vol. 15, No. 1 Ticks 183

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Agric. Sta. 66:49-55. text. ticks I. Internal Exp. Rep. ( Japanese 1959. Morphology of yato-byo Enghsh summar)'. structure of Haemaphysulis flava Neumann, 1897, Namba, N. 1958. Ecological studies on Haemaphy- Female. Ann. Rep. — Ohara Hospital, No. 6. salis hispinosa. a liarmful tick in the pasture of 3(1):8-18. (Japanese text. Engli.sh summary.) northern Japan. Hokkaido Nat. Agric. Exp. Sta. Ohara, S., and M. Sato. 19.53. Tick-borne infection Rep. 50:1-99. (Japanese text. English summary.) of yato-byo. Rinshoigaku 8:87. N'.AMBA, N. 1963a. On methods of collecting unfed Ohara, S. and S. Tamura. 1958. A case report of ticks on vegetation in grazing pastures. Jap. Soc. a tick infesting die hirnian anus. Bull. West Br. 9:21-28. text. 14:23-24. text, Grassland Sci. J. (Japanese English Jap. Sanit. Zool. (Japanese mimeo- summ;u-y. graphed.) Namba, N. 1963b. Researches on the distribution Oken, L. 1818. Sogenannte giftige Wanze in Per- of ticks and their habitat in pastures of Hokkaido. sien. Isis. p. 1567-1570, PI. 19, Fig. lA. 80:103- Oliver, Res. Bull. Hokkiiido Nat. Agric. Exp. Sta. J. H. Jr., and C. Bremner. 1968. Cyto- 114. (Japanese text. English summiin. genetics of ticks (Acari: Ixodoidea) 3. Chromo- Neum.\nn, L. G. 1897. Revision de la f;unille des somes and sex determination in some Australian i.xodides. 2e memoire. Mem. Soo. Zool. Fr. 10(3-4): hard ticks (I.xodidae). Ann. Ent. Soc. Amer. 61 324-420. (4):8.37-844.

Neumann, L. G. 1899. Revision de la famille de.s Oliver, J. H., and H. Tanaka. 1968. Personal com- ixodides. 3e memoire. Mem. Soc. Zool. Fr. 12:107- munication. 294. Olsufjev, N. G., and V. G. Petrov. 1960. Discovery Neumann, L. G. 1901. Revision de la famille des of HacmuphysuUs concinnn Koch ticks naturally Lxodides. 4e memoire. Mem. Soc. Zool. Fr. 14 infected by the tularemia agent. Trud. Inst. Zool. (2-3): 249-372. Akad. Nauk. Kazakh. USSR. (12):54:56.

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184 Bkk;ham Young University Science Bulletin

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bats {Piptsiri'llux ahnwnis) collec-tcd in Niigata tlxodes uriae ( White) =;)iif».i ( Pick.-Cambr. ).

City, Japan. Niigata Igakkai Zasshi 69(1):7-12. Zool. .\nzeiger 12,3( 1-2) : 12-17. Japanese te.xt. Sharif, M. 1928. A revision of the Indian I.xodidae ( Saito, Y. 1958. On (he intenial organs ol lanal, with special reference to the collection in the nymphal, and adult stages of the tick, Ixodes Indimi Museum. Rec. Indian Mus. .30( 3) :217-334. perxulcutu.'i. Sanit. 9:70. transmitted persulcatus Jap. J. Zool. Sharif', M. 1938. Diseases by the Indian Japanese te.xt. species of ticks and the possibility of their preven- ( Saito, Y. 1959a. Studies on the transovarial trans- tion through biological control. Ind. J. Vet. Sci. mission of Pasteurclhi ttilurcnsis in the three tick 8(4):3.53-.366. (reprinted 1939.)

species ( Ixodes pcr\ulviitu\ persulcatus, Ixodes Shioamori, M., K. Aso. and A. Yajima. 1953. On jluvti). in Prefecture, japotierisis. and Uaemaplujstdis Jap. J. Bad. the Ixodidae Kumamoto Kvushu, 14:565. (Japanese te.vt.) Japan. Nihon Juigakkai Zasshi (Jap. |. \'et. Sci.)

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rearing and life history of three tick species ( Hac- cow 40(12):51-56. ni/ipliiisalis flaia. Ixodes persulcutus persulcatus, SuGiMOTo, M. 1935. Note on the Formosan ticks,

and Ixodes japoncitMs) in Japan (Acarina: Lxodi- 1. Taiwan no Chikusan 3(9):Suppl. 1-33, 3 PLs. dae). Acta. Med. Biol. 7(4):303-321. (Japanese text.) Saito, Y. 1960b. Studies on Lxodid ticks, IV. The SuGiMOTO, M. 1936a. Note on the Formosan ticks, internal anatomv in each stage of Hdemapht/salis II. Taiwan no Chikusan 4(1):1-11. (Japanese flava Neumann', 1897. Acta. Med. Biol. 8(3): text. 189-293. SuciMOTO, M. 1936b. Note on the Formosan ticks, Saito, Y. 1962a. Studies on ixodid ticks. VI. Addi- IV. Taiwan no Chikusan 4(.3):l-20. (Japanese tional report of tick rearing, with a presentation text. of a new tick container. Acta. Med. Biol. 10(2): SuGiMOTO, M. 1936c. Note on the Formosan ticks.

Ilhis. Soc. Vet. ) :577-.590. 127-146. J. Cent. Med. 49(7 (Japan- Saito, Y. 19621). Isolation of Pasteurclhi tulurensis ese text. from ticks chiggers panusitic on tularemia- SuGiMOTO. M. I9.36d. On a new v.irietx and ( Haema-

wild hares, with some tularemia transmission ex- phi/salis cornif^eru \ar. taiwana n. var. ) and a

periment. Acta. Med. Biol. 10(2) : 147-159. Illus. new species ( Ixodes taiwanensis n. sp. ) of ticks Saito, Y. 1964. VII. animals. Studies on ixodid ticks, Notes from the Formosan domesticated J. Soc.

on the ticks infesting badgers in Japan with a Trop. Agric. Taiwan 8(4) ;336-346. ( [ap.mese description of Ixodes tauuki n. sp. Acta. Med. Biol. text. 12(l):59-66. SuciMOTO, M. 1937a. Notes on the ticks in the Saito, Y. 1967. Studies on ixodid ticks, IX. Ixodes Formosan mountain reservation for the Aborigines. inotuispiuosus u. :.30.3-,3.3.5. sp. from man in Niigata Pre- J. Cent. Soc. Vet. Med. .50(5) (Japanese fecture, Japan ( Ixodoidea, I.xodidae). Acta. Med. text. Biol. 15(2): 107-112, Fig. 9. SuciMOTO, M. 1937b. Notes on the ticks in Kvushu

Saito. \. 1969. .\. and islands. Soc. Studies on ixodid ticks, Ilacnui- Okinawa J. Cent. Vet. Med. 50 pht/salis me^aspinosa n. sp. ( Ix(xloidea, Ixodidae) (9):597-615. (Japanese text.) from Kanagawa Prefecture, |apan. Acta. Med. SuciMOTO, M. 19.37e. On the uvmph and larxa of Biol. 17( 1 )':87-96. And>lijomma testudinuriuin Koch. 1844. Trans. Nat. Saito, Y., M. Kuhota, A. Y.^jima, T. Wat.^nahe. .\nd Hist. Soc. Formosa 27:160. K. Kamino. 1965. Studies on ixodid ticks, VIII. SuciMOTO, M. 19.37d. Studies on the Formosan ticks On Haemaphijsalis hi.spinosa Neumann, 1897 in (9th Report). (On the tick, Haeniaphi/salis uishi- Niigata Prefecture. ijatnai Sugimoto, Japan, with some supplemen- 1935). J. Soc. Trop. Agric. Tai- tary observation on bovine piroplasmosis. Acta. wan 9:287-290. (Japanese text. English summary.) Med. Biol. 13(2): 143-159. Illus. Sui'iNO, F. 1897. Nuovi Ixodes della Birmania. (Nota Saito, Y., and S. Ohara. U)61. Studies ou ixodid Preventiva). Atti. Soc. Veneto-Trent. di Sci. Nat. ticks. V. Further studies on the reactions of the 2. Ill, f:i.sc. 1:2,30-2.38. skin of laboratorx animals to the bites of immature Suzuki, T. 1930 Notes on the male of a tick, Ixo- ticks. Acta. Med. Biol. 9(l):l-32. des affiuis. I.ansani.i 2(16):90-92. (Japanese Saito, Y. S. Ohara, and T. Unac.-vmi. 1960. Studies text. on ixodid ticks. III. Tanaka. H., K. Sato, Comparative observation on Y. Sugita, J. Okabe, Y. Saka- the histological changes of host tissue caiised b\ GAMI. T. CiIIKATSUNE, AND T. YoSHIDA. 1960. tick bite. Acta. Bed. Biol. 7(4)323-329. On a tick para.sitizing the human perianal region. Sautet, J. 19.36. Invasion domicilaire de Ehipiceph- J. Osaka City Med. Cen. 9(3) :743-744. (Japanese alus saufiuineus et de Teutana truinKumamoto Prefecture. Cuinea. Z. Parasitenk. 6(3) :416-431 J. Cent. Soc. Vet. Med. 24:6-12, 25(3). (Japanese ScHiL/.E. P. 1935. Zur Zeckenf:uina Formosas. Zool. text.) )

Biological Series, Vol. 15, No. 1 Ticks 185

TouM.\NOFF. C. 1944. Les Tujue.s ( I.xodoidea ) de White, A. 1852. Appendix. Journal of voyage in rindtxliine. Institute Pasteur de I'lndochine, Sai- Baffin's Bay, by P. C. Sutherland, 2:210. gon, 220 p. Wilson, S. V,. 1946. Sea.sonal occurrence of Lxodi- Trapido, H. 196.5. Notes on eritical Asian Hacma- dae on cattle in Northern Province. Nyasaland. physalis speeimens in Euro{>ean museum collec- Para-sit. 37:118-125. tions, with designations of lectotypes and a neotype Yajlma, a. 1942. On the Ixodidae collected from I.xodidae). Proc. Soc. W;Lsh. Assn. (Acarina: Ent. 67: Iwate and Aomori prefectures. J. Anim. 152-165. Hyg. 10(6):499-510. (Japanese te.xt.) TsucHiE, v., A. IsniK.wvA, AND M. Okui. 1966. Stud- Yajima, a. 1950. On tlie forms of female and each ies on ticks in pasture (II), (III). Report of the stage of Haenuiplu/S(dis ias Kishida. 1936. Jueki Institute of .\ninial Health of Shimane Prefecture. Chosasho Kenkyu Hokoku 25:197-200.

13:1-15. (Japanese text, mimeographed.) Yajima, A. 1955. On two new species of ticks ( Ixo-

UcHiKAWA, K., AND A. Sato. 1968. Tarsal cliaetota.w didae) from Iwate Prefecture. Jap. J. Sanit. Zool. of Argus japomcus Yamaguti, Clifford, and Tipton, :52-53. Japanese text. 6(1) (

1968. (Ixodoidea: Argasidae). Jap. J. Sanit. Zool. Yajima, A. 1963. On the distribution of the i.xodid 19(3):157-161. ticks on the cattle in the pastures in Japan. Jap. UcHiK.AWA, K., A. S.\TO. 1969. cK'currence Sanit. Zool. 103. text.) AND The J. 14(2): (Japanese lividus Tipton. of Argus japonicus and Ixodes in Nagano Yamaguti, N., C. M. Clifford, and V. J. Pref., species, as- Japan. J. Med. Ent. 6(l):9,5-97. 1968. Argas (Argas) japonicus new

I'cHiKAWA, K . .\. Sato, and M. Kucimoto. 1967. sociated with swallows in Japan and Korea ( Ixo- :4.53-459. Studies on the argasid infesting the Japanese doidea, Arga.sidae). J. Med. Ent. 5(4) iirhica. N., the House Martin, Dclichon Med. J. Shinshu Yamaguti, and S. Inatomi. 1961. On mor- Univ. 12(2):141-1.55. phologN' of larva and nymph of Argas species col- W'ARHrBTON, C. 1908. On some new and obscure lected in Okayama Prefecture, and some additional its species of tlie genus Hacmaphystdis of the Ixodi- observations on biology. Jap. J. Sanit. Zool. dae. Proc. Cambr. Philos. Soc. 14:508-519. 12(2): 142. (Japiuie.se text.) \\'.\rburton, C. 1913. On four new species and two ZuMPT, F. 1940. The Rhipiccphalus species of the new \arieties of the ixodid genus Hncmaphysalis. USSR. An account of variabilitv in the sanguineus

Parasit. 6(2) : 121-1.30. group. Z. Parasitenk. 11:400-409. W'ahhurton, C. 1932. On five new species of ticks ZuMPT. F. 19,50. Preliminary sttidv to a revision of (Arachnida, Ixodoidea). Parasit. 24:.5.58-,568. the geniLs Rhipiccphalus Koch. Docum Mocambiciue Warburton, C, and G. H. G. Nuttall. 1909. On 60:.57- 125. new species of Ixodidae with a note on abnormali- ZuMPT, F. 19.52. The ticks of sea birds. Australian ties observed in ticks. Parasit. 2:.57-76. Natl. Antarctic Res. Expedition Rep. Series B. 1:12-20.

APPENDIX 1

Notations on the Literature Cited

J. Jap. Soc. Vet. Sci. 16. 1-10, Yajima's (1956) list was often referred to as a source of 1-11. 1936, and in the pp. of contains two papers information by Keegan and Toshioka (1957). Since it is a 1937. 1 iirthermore, the paper 1937a in Taiwan no Chikusan mimeographed note and has not been published yet, it is (with the same title) published (1937e) is excluded in this paper. 5(3):l-25, 1937 and 5(4):1-15, 1937. His paper Serial reports by Sugimoto entitled "'Note on the also found in J. Jap. Soc. Vet. Sci. 16( 1), pp. 11-16. 4" ticks domestic animals in I ormosan ticks, part 1, 2, 3, were published in Taiwan no The paper "Studies on the of the

( is an ( hikusan .3(9). 1935,4(1), 1936,4(2), 1936, 4(3), 1936, re- the Far East" by Itagaki. Noda, and Yamaguchi 1959) spectively, and these four papers were at the same time English translation of their 1944 paper, and the contents of published in Bull. No. 118, 1936, Dcpt. Agric. Govt. Res. both are identical, except that H. jezoensis is synonymized of localities is Inst., 1 orinosa, Japan. Likewise, another of his papers under H. japunica and spelling Korean

1 1 so as to be up to date in the later (1959) publica- < 1936d) appeared both in the Taiwan no Chikusan 4( ), pp. amended tion. 186 Bkigham Young University Science Bulletin

APPENDIX 2

'° Collection Records

Species Host or llabitjt Area Preli'Ltutc Locjluy Collection or Literature Data

Argas japonicus Di'lichon urhica dasypus Japan Nagano Agcmatsu-cho *66-J-0032.Coll. Toshioka. 17 Jul •66(42 (House Martin) I .•»4 M. 227 N, 39 I. 2 F. 2M. 21).

'66 Matsuinoto-shi Coll Vamaguli, 7 Jul (5 1 . 5 M. ION. 10 L).

'66 Coll. Sato. 1 3 Sep (5 1 , 5 M, 5 N).

Iwatc lwai/-umi-cho #66-J-0()43.Coll Yaniagud. 7 Aug •66. IWH I , 17H M. 130 N.J/-.

2 M. : 1 1

Tt)chigi Chu/enji *66-J-()()47.Coll Hatovanui. 24 Aug '66. (ION).

Tokyo lacliikawa M H C:66-1-II04S. coil, lloluhcc & Sugl-

:ili yania. 24 Jun '66. (36 I . M. 153 N. 25 L, J/-'. 2M)

Shiniane Iwannod.i-shi #66-J-O()50. Coll. Holubec & Sugi-

yaina. 7 0ct'66. (16 I . 11 M.47 N).

! ukushinij Shinohu- r.ika> u »6)(-J-(l336. Coll, Suzuki & Sasa-

gawa. 10 Jul '6S. (700 I . 600 M. 1.100 N),

Nakanosawa, #6X-J-0337. Coll. Suzuki & Sasa- Inawashiro gawa. 12 Jur6X.(4 N).

llirttndo daurua japontca Oka>ania #66-J-()(l37. Coll. Mizusawa & ^66. I Striated Swallow! llatoyama. 21 Sep (89 1 . 57 M. 108 N. 34 L.J/-', 2M.2 I.).

^60 Coll, Yainaguti. Sep. (5 I . 3 M.4 N. 7 L).

Totlori Yonago-shi #66-J-()()53. Coll, Mizusawa &

Haloyama. 1 9 Sep '66. (45 1 .

62 M,89 N.J /-, J.W).

Miyazaki Takachjho-cho #66-J-()044.Coll, roshioka&

Suzuki. 13 Scp^66. (63 V . 30 M. 496 N. 5 L.J/-. J.Ul,

Gifu Mitakc, Kani-gun ^66-J-()()62. Coll, Tanaka. IK Oct

'66. I I Nl.

Okayama Nnnii-shi Inatomi* "iamaguti. 1960:17. Yamaguli & Inalomi. 1961:142 Coll. "60. Yaniagull. Sep (I . M. N. Ll.

Dcliihon urhua dawpus Nagano Agcmatsu-cho Har.i. 196.t. (oil. Ilara i^\ pirsitiis). (House Martin)

Matsunioto-shi. Uchikawa. 1969:96. Coll.

Hakuha-mura. llchikawa. 20 Mar '67 - 7 Apr '68.

Ageinatsii-cho. (A. N. 1 ). Akasliin.i-clio.

1 ukusliinia-cho. Hotaka-etio. lida- shi. Nagano-shi. Okaya-shi. Otari- niura. Saku-shi. Sakai-niura & Ueda-shi

Sleeping tpiarters Korea Kyongsang- Taegu YaniaguO. Clifford & Tipton. 1968: pukto 458. Coll. Tibbets. 20 May '53. (51.1 M. 3 N).

:'(1 iiiituhcrs in iwiientlicscs inilicale numlH-r of spt'< inions sent In Mr (.len .M Knliis fni i.li'iilifu.itiiin Biological Seuiies, \'ol. 15, No. 1 Ticks 187

Argas japoiticus Swaiinw nt-sl Korea Kyonirvinj;- Shin C lion l)i pukto

Argas Vesperiilio \upiraiut^ lj|Mn I ukusliiin.i Slidji!, Bango-cho icsficrfilHinis (Irostcd Bat)

Pipisrrcllus ahramus (Hal)

Ornithodoros

i-api'iisis 188 Brigham Young University Science Bulletin

Amblyomma Cyclemys flavomarginata Ryukyu Gosj-dukc, ^66-R-OOI7.('oll. Mizusawa, 19

geoemydae flavomarginata (Turtle) Islands Iriomolc (Iv) Jun "66.)/ h'. 10 \. 14 I ).

Land torioisc Okinawa -ft6-R-U006. Coll. Pennington.

Oct 'bt.{4 N. 2S I.).

Gcoeniyda spcngleri ^ una. Okinawa =6N-R-(I()15. Coll Mi/iisawa. Apr japomca (Turtle) "68. (11 N).

Clemmys japoniva Japan Kagoshinia Kunijranii. Tane- Keegan & Toshioka. !957:S. (as (Turtle) gasl ima A. malavanum) Coll. Toshioka. 19

Jur55 (1 1 . 1 M, I N).

Cyclcrriys flavomarginata K\ ijk> u I'utanaka. Ishi- Kecgan & Toshioka. 1957:8. (as flavomarginata (Turtle) Islands gaki (Is.) .1. malavanum) Coll, Keegan &

loshioka. 6 Nov "55. (I I . 1 M).

Hanoshiro. Ishi- Keegan & Toshioka. 1957:8. (as

(Is .1 gaki I malavanum) ("oil. Keegan &

loshioka. 27 0e( *55. ( 1 Nl.

Gosa-dake. Iriu- Kaw.ishima. 1963:103. las,l, mote (Is.) malavanum) ("oil. Kawushima. 14

Aug'62. (9I-.4 M. 3N. 1 L).

Keegan & Toshioka. 1957:8. (as A. malavanum) Coll. Keegan & Toshioka. Nov '55. 406 MCil -IHI. ll N)

(icocmyda spcuglcn Yona. Okl^.l\^a K.iwjshniia. 1963:103. (as.-l. (Turtle) malavanum) Coll. Kawashhiia, 6 Jul "62.(3 N. n L).

Turtle West ot Arakawa. Keegan & Toshioka. 1957:8. (as^l. Okinawa malavanum) Coll. Hubert. IS May '55. 406 MtU -25. (6 N).

Man Mountain near Keegan & Toshmka. 1957:8. (as.4.

Ishigaki ( ity. malavanum) CoW Keegan & Toshioka

Islugaki (Is.). 27 Get "55. ( I N). (Probably a stray troni the turtle ot 27 Oct '55 eoUection examined the same day).

Amblyomma l.aticauda scmifa- Khigaki (Is I -66-R-(XH)8. Coll, Toshioka & nitidum sciata (Sea snake) Keegan. 10 Nov '62. (/ /')-

^66-R-nf)09. Jun '57. (/ F)

A mblyomma Sus scrofa leucomysiax Japan loehigi =6(( J-045.(oll Aki>ama. Nov testudmarium (Wild pig) '60.1/ h\ n.M)

Ryuk\ u Kagoshima .Ashiken. Anianii #67-J-0246. Coll. Mizusawa. 20

Islands Oshiina Sep '67. (2 I . I Af)-

\'u\van. Aiiuiini -=^67-J-0256. Coll. Mi/usjwa. 1

/-". Oshinia Oct "67.(3 I . / / M).

^67-J-0272.ColI Mi/usawa, Apr

'67. ( 1 M).

#6«-J-()310. Coil Mi/usuwa. 15 Mar 'bH.{7 F).

*68-J-0318. Coll. Mi/usawa, 1 1

Apr '68. (2 I . 2 M, 4N.5 L.

3 N. 5 I. ).

Ashiken. Anianii ^670-0271. Coll. Mi/usjwa. 19 Oshima Sep *67. (/ ;V).

Islngaki (Is.j ^55-R-()17. Coll. Keegan & loshioka. 4 Nov '55.(7 F. 5 /.),

Domestie pig 'I'uvK.in, Am.inii ^65-J-()OOI, Coll. Hajime, 13 Dee OshlMKl •65.(J/-", J M) Biological Series, Vol. 15, No. 1 Ticks 189

Amblyomma DomestiL- pig Ryukyu Kagoshimj Yuwan. .Anianu #65-J-0002. Coll. Ila)nne. Nov '65, Isbnds Ifstudmanum Oshima I / \U.

Ukenson. Atiiami #67-J-0067. Coll. I ipton & Mizu- Oshima sawa, 18 Mai '67.(1 h. 12h L.

5 F. 5M. 19 1.).

#67-J-()073, Coll, Tipton & Mizu- sawa. 19 Mar'67.(/ F. .^M).

#()7-J-0077.C'oll. I ipton & Mi/u-

sawa. 1,1 Mar '67. (//•'. / A/).

Cenus nippon Ibaraki #67-J-OI83. Coll. Yamaguti. 22 (Sika Deer) Mur'67,(/Af).

Shizuoka Misakubo-cho #67-J-0293. Coll, Su/.uki, 4 Dec '67, (/yW).

Domestic dog Ryiikvu Ragoshiina \'u\van, Amami #67-J-0273. Coll, Mi/usawa. Apr Islands Oshima '67.1/ F).

Penialagiis Jurnessi „ „ Ukenson. .Amami #67-J-00b9. Coll, Ttpton & Mizu- (Ryukyu Rabbit) Oshima sawa. 24 Mar "67.(1 .16 L. 1(17 I.).

#67-J-0081, Coll. Hajnne. 1(1 F eb '67. (7 95/.).

Yuwan . AmamI #68-J-0314. Coll, Mizusawa. 5

Oshima Apr '68. ( / An.

Vegetation LIkenson, Aniami #67-J-(J082. Coll, Tipton & Mizu- Oshima sawa. 23 Mar '67. (/ /.).

#67-J-0076, Coll. Tipton & Mizu- sawa, 23 Mar '67. (97i).

#67-J-0084. Coll. Tipton & Mizu-

sawa, 24 Mar '67. (45 I.).

Yuwan. Amami *68-J-0316. Coll, Mizusawa. 5 Apr Oshima '68. (JFl-

#68-J0317.Coll. Mizusawa. 10 Apr '68, 1/ .V. ,5/.).

Japan #66-J-0031.Coll, Ucno. 28 Jul '5 7. (IF).

Man Hyogo

«•

Horse

Wild pig

Vegetation

Domestic pig

Microhyla fissipes (= M. ornata) (Frog)

Wild pig 190 Bhicham Young University Science Bulletin

Amblyomma Wild pip Ryukyu Iriomotc (Is.) Sugimoto, 1937h:612-613. Coll. testudinarium Islands Ohasi. Mar '37, II M).

Green trog Ishipaki (Is.) Sugimoto. 1937b:6l2-613. Coll. Iwasaki, (1 L).

Japan Miyazaki. Nakamura & Yajima. 1937:174. Hyogo

Horse Miya/aki Takanabc Robinson. 19:6:257. Jun & Oit '12 IN 2909a).

Ryukyu Kagushima Anuinu Oshniia Robinson, 1926:257. Jun & Oil Islands •|2(N 2919).

Boophilus Domestic cattle Japan Nagasaki CJonoura. Iki #67-J-lllll3. Coll. lipton 4i Min/u-

microplus (Is.) sawa, 2 May '67, 16 1 i.

Hatsuyama, Iki #67-J-0107, Coll. Tiplon & Mizu- (Is.) sawa, 4 May "67, {3 M. J .V).

Iwa/ato. 1 suvhiina *67-J-0l IS. Coll. Tipton* Mizu-

sawa, 6 May '67, 1 /.^ F. 7 M. I J ,V).

KagitsliiiiKi k liinan #67-J.|li:4. ( .111 1 iploii i Mi/u- sawa. Ill May '67. IK I, J/-,

.5 M. 4 ,V).

#67-J-OI 25, Coil, Tiplon & Mizu-

sawa. 10 May "67, 1 1 I, 5 F. 4 .V).

#67-J-OI26.Coll hptonii Mi/u-

/', sawa. 10 May "67,17 I , // : M I .V).

#67-J-l)l 28, Coll. liplon & Mi/u-

"67, .V/-', sawa, 10 Mav I 12 I-,

#67-J-l)l2M.( oil. lipton & Mi/.u-

sawa, HI Ma\ '67. 14 I . h /).

W.7-J-III 31, Coll. Tipton & Mizu-

/•', sa»a. 10Ma> '67,119 I . 7 71/. 4 \\

#671111 32, (oil. I ipton & Mizu- sawa, II) Ma\ '67, lA'F, 4 M).

#67J-ni 34, Coll. lipton &. Mi/u- sjw.i. III M.n '67,15 1. :i /', SM, 4 \)

#67-J-0135, Coll. Tipton* Mizu- sawa, 10 May '67,(2 V.jh: JM. 4N).

#67-J-0 137, Coll. Tipton* Mizu-

sawa, 10 May 'bl. iiO l . 25 F. V M. / ,V1.

Sakura Jinia ??67-J-013»(, Coll. Mizusawa, 14 '67, May i \\ . 4 F. I M].

#67-J-014ll. ( oil Mi/iis.iwa, 14 May '67, 1/ F)

#67-J-OI4l,Coll. Mi/usawa, 14 May '67,(/7K 5 M).

J?67-J-014:. Coll Ml/lls.iw,i. 14 '67, Mav |7 I , I M, I N, J F. 1(1 V. I \).

Ryukyu ,Sunii>o, AiiKimi -67-J-()064, Coll. Tiplon & Mizu-

Islands Oshinia sawa, 22 Mar '67,124 I ...'() /•, -V.AH.

Burcn, .Amaini =67-J-0252, Coll. Mi/usawa, 2X

/(;/'. t, Oshiina Sep '67, I 15 I , 3 M. M).

Yuwan. .Xniami .=67-J-026l, Coil, Mizusawa, 3 Oct Osliillla '67,(93 1-, 3M, 3N). Biological Series, Vol. 15, N( Ticks 191

Hoiiphiiiis DonicslK caltic Mivara. Khigaki =h(.-R-(l(l(i:. (nil, HnlulK'c & Don- tiikroplii\ Inlands (Is.) ccllc, 2 Nov '(i(>. (,V7/-', l,S At).

=?66-R-()00.1, Coll. Doucette & Su- glyama. 2 Nov '(id. (s'.s /, _\S A/).

#f.6-R-0004. Coll. Doucette & Ho- luhec, 2 Nov '66, (^.5 F. 4.i AT).

Hirai. Ishigaki -hV-R-OOl l.Coll Tiplon. K I eh

(Is.) '67. ,s (15 1 . M. 2 N)

#67-R-()IILl.C oil. 1 ipton. (. May "67, (X I . i Ml.

Korea Kurnik. Cheju Do *h7-K-lll)19. Coll. Parsons, ft May '(.7.1/ ,V|.

Cluingihont;- Macng-dong Myi>n #h7-K-()029. Coll. Cantello & Town-

pukto send, 12 Mav '67. (9 N. / F. / M.

I .V).

Uni-Song Klin #(>7-K-(10.V). Coll. Cantello & I own-

'67. .^ send. 12 May I I . 9 N. / M).

#67-K-(X)3 1 . Coll. Cantello & Town- '(.7. send. 12 Mav 1 2 i: 7 N. / K / ,Vi.

Huhalus hu halls R> uk\ 11 Hir.ii. Ishigaki #h7-R-(l(ni), C oil. Tipton, 9 I eh

(.^siatK WaUT Hullalo) Islands (is) '67, I ll 1 . 9 M, 2S N).

#67-R-0(ll2, Coll. Tiplon. S I ch

'67.(1 I 1.

Donicsdi. ^hccp Korea Isodong. Cheju Do #67-K-(l(122. Coll, Unggers, 6 May '67. (/ /).

Hoitphillis spp. Cattle Japan

Cattle, water buttalo. horse, deer, pheasant

Cattle

Horse and duniesiic cattle

Horse, cattle, domestic

sheep. ,'\ sialic Water Buttalo .

192 Hhicham Young University Science Bulletin

Cholla- Boophilus spp. Horse, cattle. domesiiL Korea Itapaki. Noda & Yamaguehi. 1944. sheep. Asiatic Water nanido 33-3'J, 1959:41 (usfi. cauJatus). BuITjIo

Cattle and horse Jupan Kuinainoto Shigemori. Aso (S: Vajima, 195.V 290-293. (as Vroboophilus cQudatus and Palpohoophilus brachyuhs).

Cattle, horse and water Ryukyu Ishigaki ( Is.), Keegan&'Ioshioka. 1957:10. butTalo Islands Inomote (Is.), las Boophilus niicropliis). Yonakiini (Is.). Matenima, Kobarna Korea K\ongii-do Seoul

Cattle Japan lokyo Nultall & Warburton. 1915:433. (as Hoopfulus sp.. tound with

//. hisponosa = H. longicornis. 16 Oct "11. submitted by Dr. Miyajima).

Horse Miyazaki Neumann. 1897:413. (as Rhipiccphalus atmulatus cauJatus) (23 r, 2 M).

Dermaccntor spp Wild pig Chiba Kiyosumi-vama Kishida. 1922a: 847-851. Coll. Aoki. 30 Oct '09 (as auralus auraiwi) (1-, M).

Domestic pig Tokyo Kishida. 1922a:847-851. (as rclicularus rcticulatus) (I-. M).

Korea Kishida. 1922a:847-85 1. (as

rcticulatus reluulatus) (I . M).

Keigen-gun. N. Kishida, 1936:142. las/;. = Kankyo ( Hamgyong- vancgaius). .According to Kishi- pukto) da. Mr. Asakawa recorded this species.

Horsi Japan Iwute Taniura, Tayama- Nakanuiraii Vajinui. 1937:162-164.

mura, Johoji- {dsreticulalus) li . M). mura

Terada. h^atc•gun. Yajima. 1942:499-510. (as Okunakayania. Ninohc- rcticulatus) (1. M).) gun. Johoji-mura, Ninohe- gun. Arasawa. Ninohc-gun.

I utaniata Pasture.

Sannohe-gun Yajima. I 942:499-5 I 0. (as Kamikita-gun rcticulatus) (W M). Shimokita-gun etc.

Tcmmabayashi- Itagaki. Noda & Yamaguchi. 1944. mura, Kamikita- 1959. (;is rcticulatus) (28 1-. giin 16 M) Aug& Sep '41 & Oct '42.

Iwatc Takizawa. Iwale- Itagaki, Noda & Yamaguchi. 1944. gun 1959. ids rcticulatus) (12 l. 10 MlOct '41.

Korea Hanig\ *uig- Unggi. Kyonghung Itagaki. Noda & Yamaguchi. 1944. piiklu 1959. as (-0^/^(61 I. 34 M) Jun *41 & 29 May-5 Jun '42. Ori- ginal description oi corcus. no indication ot type.

0- Selcnarctos tiuhctanus Japan Shi/uoka Oshinia-inura, Keegan. & Toshioka. 1 957 : 1 M japonicus Sunto-gun Coll. Asanuma.(2 M). (labelled as (Asiatic Black Bear) auratus).

Wild pig Kyoto Shi/uhara Keegan & Toshioka. 1957:10-11. Coll. Akiyama. 12 Dee '55. 406 MGL -101, closely resembling the above. Illustrated in pi. 14 ot K &. T.. reproduced. (2 M).

Kccgan & Toshioka. 1957:10-11.

Coll. Toshioka & Akiyama. 29 I eb '5h. 406 MGL =194.(2 Ml. Biological Series, Vol, 15, No. 1 Ticks 193

Dcrrriaantor sp[v Wild pig Japan Kyoto Kyoto-shi

Horse Aoniori

Iwate

Okunakavama

Korea Yuki ( = Unggi)

Hacniaphysalis Man Japan Hyogo cairifyantilala

Domestic dog

Niigata

Hokkaido

Tokyo Chiba Niigata Saitama Kanagawa Shizuoka Mic Osaka Hyogo Kochi Miyazaki

Korea Kyongii-do

Domestic cattle and Japan Nagasaki horse Niigata Ciunima Ishikawa Yam ague hi

Korea Hamgyong- pukto Kangwang- do

Japan Kumainolo

Uog Kanugawa 194 Bhicham ^'(>UNc University Science Bulletin

llafmaphysalis Dog Japan lokyo Nuttall& Warburton, 1915.491-492.

campamtlata Coll. Janson. I X94, ( *2XK6) (I A).

Nuttall & Warburlon. 1915;491-492. 18 Jun "14. submitlfd by Dr. Miyajima(#299l) d A).

Nuttall & Warburton. 19 l.^;49l•492.

1 I Sep '12. subniittfd by Or. Miyajima(*29l7l (I Al.

Hat- maphy sails Uokkaidu Ncinuro-shi, Kitaoka, May '64. {I h. J M). cominna Neniuro

Donicstii' cattle Alsukeshi. Kitaoka. Jul '6h, (/ h'). Kusluro

DonicstK cattk' and Tokachi. Kushiro, Ogura& lakada. I 927;2ll2-2(>4. horse Ncmuro & Kitami

Korea

Horse

Korea

Hamgyong- Unggi pukto

Japan

Dog Saga Biological Series, Vol. 15, No. 1 Ticks 195

Hac maphy sails Phasianus cokhicus Japan Niigata Shukunegi. Sado

JoeniCi rohusfipcs ( Pheasant) (Is.)

Unie/u. Sado (Is.)

Vegetation

Pheavint Kagoshinia Akune

Niigata

Hacmaphysali^ Horse Aomori Towada flava

Dog Tochigi Kuzuu

Mie

Towada

Kanagawa Yamakita-cho

Fukushima I ukushima-shi

Iwate Ohinado

Kanagawa Miyagasc, Kiyo- Kawamura

Yamanashi Nohara, Doshi- tnura

Nycterciitcs procyonottjes Miyazaki Tomisaki liierrinus (Raccoon Dog)

Vulpes v'ulpcs japomca Kyoto Hanase-mura (Red I-o\»

Sus scrofa leucomystax (Wild pig)

Iharaki

Aichi

Ccn'us nippon (Sika Deer)

Domestic cattle

Vegetation 196 BntCHAM Young University Science Bulletin

Haemaphysalh I.epus hrachvunts Japan Kagoshiina Ogawa. Kokutni-' ^68-J-l)339, Coll. Yaniamoto. 16 jlava angxiHtidi'fis Jan-.l Mar '68, (9 I. 42 M. 18 N. (Hare) 6L).

Horse Iwate Okunakayaina (No. 25.'t5), Coll. Watanabe. 16

VIII 1939.(1 I", labelled as //. waianahei n. sp.).

Dog. liorsc. kow. hare, Tokyo? Neumann. 1897:333-336.(11 I",

vegelalion Saga-' 7 M. I 3 N. L - type scries-), Miyagi? presented by Hrot. Suto at Tokyo, Yariiagiiehi at Saga, Miskina at Sendai. Neumann states nymphs were from hare & cow, larvae from eow.

Horse, vegetation, Neumann. 1905:238, 141 1:1 12. (as Lcpus sp, (Hare) flaia var. annata oi jlava armata) cited by N. & W., 1915.

Cattle, horse, dog Hiroshima Donit/, 1905:105-134. Donitz Shiniane expressed doubt con<.crning

Kyoto.' I aniba Neumann's idcntitiLation tor the specimens m Herhn Museum.

VVarburtun, 191)8:510-512. Same specimens as N. & W.. 1915. Piesented by Donit/..

Dog Mic isG NuttailA Warburton. 1915:408-

410.(1 I . 1 M).N. 893. Also cited Neumann's record (1911).

Cattle or horse Hiroshima

Wild pig Chiba Kiyosumiyama, Kishida. l922a:852-853. Coli.

Awa-gun Aoki, 30 Oct '09.(1 M)(as//. /.).

Dog. horse, cattle, Kishida. 1927:985. wild boar, serow

Korea

Dog, cattle & horse

Cattle and horse

Cattle & dog

Horse and cattle

Horse

Horse and dog

31 species hsled

Domestic dog. Sch'nantos thihclanus japonicus (Asiatic Hi.ick Biological Series, Vol. 15, No. 1 Ticks 197

Hacmaphysalis Hear). I.ipiis brachyurns Japan Miyagi jlava (Marc). Sciurus In Yaniagata (Squirrel). Phasianus Nagano colchictis tohkaidi (Green Kyoto Pheasant). Phasianus Saitamu soemnwrnngii scintillans Shi/uoka (Hondo Copper Pheasant)

Dog Kcegun & loshioka. 1957:14. Ineluding Holotype o\' H. orientalis Kishida, l*)34. (rom Tokyo Prcleclurc.

Japan Because o! ubiquitous distribu- (Honshu. tion, name of prclcctures and Shikoku, collecting data arc not given Kyushu) here.

Man. domestic dog, Japan Niigata Mainland and Saito. 1959h: 193-209. Coll. Nyctcrcutcs procyonoidcs Sado (Is.) Saito, "56-'59. Immature forms (Raccoon Dog). Mustt'la on small mammals and birds. sibirica itatsi (Weasel). (A. N.L). Selcnarctos tfnbclanus japonicus (Asiatic Black Bear), icpus brachyurus angitstidens (Hare), l.cpus brachyurus lyoni (Hare), Apodcmus spcciosus sado-

cnsis (1 leld mouse), Phasianus cokhicus rohustipcs (Green Pheasant), (iarrulus glandarius toku- gawac (Japanese Jay), t'rn- beriza spodocephala per- sonata (Japanese Bunting)

Cattle, l.cpus brachurus Kakudayama. Saito et al., 1965:145-146. Coll. angustidcns (Hare). Meles m. Sasagamine Pasture, Saito. II Sep '62-17 May "65. anakuma (Badger). Nye- Ohsadorynkan Pas- (I-. M. N. I), tereutcs procyonoidcs turc vivcrrinus (Raccoon Dog), Strcptopclia o. oricn-

talis { I astern Turtle- dove), Phasianus cokhicus robust ipes (Green Pheasant) and vegetation

Haemaphysalis Sus scrofa riukiuanus Ryukyu Kagoshinia Ukenson. Amami #67-J-U216,Coll. Iipton &. Mizu- formosensis (Wild pig) Islands Oshima sawa. 18 Mar "67. (/ F).

Pcnlalagus furncssi Yuwan. Amami #bS-J-()32U. Coll. Mi/usawa. 5 (Ryukyu Rabbit) Oshima Apr "68.(7 Af).

#68-J-0324. Coll. Mi/usawa. 5 Apr "68, (I N).

#68-J-0325. Coll, Mi/usawa. 5 Apr '68.(1 N).

Vegetation Ukenson. Amami #67-J-0217.Coll. Tipton & Mizu- (Jshima sawa. 23 Mar '67.(7 AD.

Yuwan. Amami #67-J-0266. Coll, Mi/usawa. 6 Oshima Oct "67,(7 m.

Kitaoka. 1970:73. (oil. H. Haemaphysalis calves Japan Shi/uoka I oot of Mt. Fuji fujisana (700 m.) near Toyota. (A. N, L). Tujinom,iya

Nov Haemaphysalis Domestic pig Yuwan, Amami #65-J-0003. Coll. Hajime. hvsrhcis Oshima '65A5F.JM).

Toshi- Sus scrofa riukiuanus Iriomote (Is.) #55-R-021. Coll. Kcegan & rWild pig) oka. 10 Nov '55. (iF. 5 A/).

Iriomotc (Is.) #55-R-022, Coll. Kcegan & Toshi- oka.5Dcc'55.(/ F. I M. I IV).

Kagoshima Ukenson, Amami #67-J-(J066. Coll. Tipton & Mi/u-

Oshima sawa. I 3-19 Mar "67. (9 i". 3 M. 9F, WM. fiX).

#67-J-0075. Coll, lipton & Mi/u- "67, (.;/'. /V). sawa. I 3- 1 9 Mar / M.l 198 Bhicham Younc Univehsity Science Bulletin

Hacmaphysalis Siis scroja riukiuamis Jjpan Kaj:oNhiinu llkcnson, Aniatni #67-J-(UI7«, Coll. Iipcon & Mizu- hyslricis (Wild pig) Oshima sawa. 13-14 Mar -ft?, (6 1-. 2 M. h F. :: M)

R>uk> ti Vainatosoii. Amaini #67-J-()07l.{ oil. Mi/usawa, 21

Islands Oshiina Mar '67.(4 1.4 M. 2(1 N. / F. 4 M. J .V).

Asiiikcn. Aniaini V67-J-024.S, Coll. Mi/.dsawa. 2(1

Osliinia Sep 7,7.(6 \ .jl F. JOM)

Yiiwaii. Aiiiami a(,7-J-ll255.('oll. Mi/usawa. 1

Osliinij Oct '67. (/ /. / 1/1.

*67-J-02.S7.(oll. Mizusawa. I Oct '67,(1 N. // h. 4M).

Ashikcii. Ainanii -67-J-027().

Dog Yliwan. AmaTiii a6.S-J-l)ll(l4. (oil llaMnic. Dec Oshinia '65.(.W-'1-

I'kcnson, Anianii ffh7-J-(l21S,( oil llajimc. 1,1 OsIiinia Aug'67.(/ AD.

#67-J-(l214. Coll. Ilajinic. 1,1 Aug '67. (2 Fl.

Yuwan. .'\ni.iiiii »67-J-(l274, Coll Mi/usawa. Apr Oshirna

#67-J-li24S. Coll. Mi/usa\w. Ill

Sep '67. ( / Fl.

#67-J-ll2h.!,Coll KiinuraAi Mi/u- '6 sawa. 4 Oil 7. ( / /I.

Pcnlala^ts furncs^i I'kenson, Am.inii #67-J-ll(16X. Coll, Tipton & Mi/u- (Ryiikyu Rabbill Oshiina sawa. 24 Mar '67.(1 N. / ,V).

Vegelution #67-J-(l(l7(). Coll. Tipton & Mizu-

sawa. 2.1 Mar '67.(1 I . 5 /V).

"Iiiwan. .^inann #67-J-0244. Coll, Mizusawa. 26 Oshiina Sep '67. (J f).

#67-J-0264, Coll. Kiniura & Mizu- sawa. 4 Oct '67. (2 An.

#67-J-027N. Coll. Kiniura & Mizu- sawa. K Oct '67, (/ /).

ff67-J-027,S. Coll, Mizusawa. Apr '57. (/.A/I.

#67-J-0.1IIX.( oil Mizusawa. 6

Oct '67.(74 I.. / /• reared).

.V»,^ scrttfa riiikiuaniis Inoda, Ishigaki Keegan & loshioka. I4,S7: 1 5, Coll,

',S.S, (Wild pigl ( Is. I Keegan & loshioka, 4 Nov

.5 (I I , M, 1 N,4 1 I,

Mountain near Keegan i loshioka. 1 457: 1 5, Coll, Inoila, Ishigaki Keegan & loshioka. 8 Nov '.S.S,

(Is.) (7 I . 17 M, 26 N. IS L),

Cow Yonakunills,) Siiginioto. 1437a 31.V 14371,604,

Coll. (Ilka. 6 Oct '36.1 I I ),

.Vi/,v scrofa nuktuanus Otoini, Irioinote kju.ishiina, 1463 1113 Coll

'62. I 1 (Wild pigl lis I Kawasliiina. IS Aug |2 . M,

2N, I 1.1.

Sus scrofa ssp. Mountains near Iloogstraalel al.. 146.S:477,

(Wild pig) Inoda, Ishigaki (2 1,4 M, 6 N. I L). (-RMI (Is.) .14123): Coll, Keegan & Toshioku.

8Nov'.S.'i.(2 M. 1 I . 2 N. 1 1) (-IIII 53261

Shrubs C liiiukj. Okinawa Moogslraal et, al,. 1465:477, (oil,

llardcaslle. 31 Aug '46. 1 1 M). JfRMl. 27531. Hioi.oGicAi. Sf.ries, Vol. 15, No. 1 Ticks 199

Hacmapbysalis (Iround Ryukyu Tanyu-yaina. Hoopstraal ct al., 1965:477. Coll. hvstricis KbtuK Okinawa Werner. 13 Jnl'51.(2 l).(MCZ).

Japan Neumann. I 9U 1:2(> l-2()2. (as hispinosa. 2 I).

Haemaphysalis Caphcornis crispus Nagano ShiiTiajuna #53-J-005. Coll. Sakaguchi. 24 laponica (Scrow) Nov'53, (/ A/).

Kanagawa Tanzawa Mis. #6K-J-l).V)4.Coli. Isucliiyj. 17 Mar "68, l2 m.

Ci'nus nippon Tochigi Kiniigawa *56-J-(l34, Coll Toshioka & Aki-

(Sika Deer) yama. 2,S I cb'.Sf.j/ A/).

OonK'stK t..ittlc Hokkaido Ktittan. lokachi #67J-()I^X. Col! jjuigichi. 13

May '67. ( 14 N,/") A, (> M).

Dog Ko/awa, Shiribeshi #67-J-()322. Coll. Tosliioka & Sasa- gawa. IS Jul '67. (/ /').

DonK'stK cattle Higashimokoto. #55-J-()64, Coll loslnok.i & Aki- Abashiri yania, 13 Aug '55. (/ M).

I.i'pus hrachvurns Nagano Asanunia. 1 95K: 274. Coll.

brachvurns (Harel Asanuiiia. '5S, ( 1 N).

Horse Keegani Toshioka. I 957: 1 7. Coll

Yajinia. May '4(1.(1 I . 1 M) (labelled as cuncinna).

Korea Kecgan & Toshioka. 1457:17.

Coll. Asanuma. Jur4i . ( I M). (labelled asjczoemis).

Hanigyung- llnggi & Kyonghung llagaki. Noda& Yamaguchi, 1444. pukto 1959.(as/czmviA/.v in 1944, japonica in 1959).

Jjpjn Nikuppu, Hidaka Memambetsu. Abashiri Abira. Yututsu. Ihuri

Cattle & horse Sapporo &. Kushiro OguraAc lakada. 1927:206. (as jczoensis).

Warburton. ! 908:5 I 2-5 I 3, (as japonmca). Numerous males. type specimens, found \\\\h flava ut the Hritish Museum (see discussion).

Dog. horse, cattle, Neumann. 1911:112. {.\^H. Jlava wild pig, Capricornis Jlava) crispus (Serow)

Capricornis crispiis Hondo Nutlall& Warburton. 1915:402-403. (Scrow) (as//, japonica) Same specimens as Warburton's types. Coll, The Duke otliedtord.

Harniaphysalis Ccr\-us nippini Mic Ouchiyjina #54-J-006. Coll, Asanuma. 17 Jan kitaokai (Sika Deer) '54, (.?/•*. JM).

#67-J-Ol7X.Coll. Yamaguti.K Mar '67.(7 Af).

#67-J-OIX5, Coll. Yamaguti. 22

Mjr'67.(6 1 .8M. /M/-'. 9 A/).

Shiga #67-J-(n80. Coll, Yjmaguti. S Mar 'blA'iF. JAf).

#66-J-OOI9,Coli. Sugiyam.1 Molu- Domestic cattle Shunane Sanbe. Oda-viii & bee. 6 Oct '66. (Aft h').

#670-0286, Coll. Yamaguli. 4 Nov

'67.(32 I-. II M,5N,.Vf. 2 M).

#67-J-0289, Coll. Yamaguti. 4 Nov '67. (//•). 200 Bhioiiam Young University Science Bulletin

1 Haemaphysatis Capriconiis crispus Jjpan Kanaga\^a . I an/d\va Mts ?f6H-J-l)3(l5,Ci)i:, Isuihiya, 5 Mar

kilaokai (Scrow) '68.(1 I ).

Caltlf Nagasaki Nakano-mura. Kita- Keegan & i'osliioka. 1957: 16. (as [iiatsuura-gun //. iiwrmis) Coll. Yajmia. 28 Jun '52.(1 I).

Dcct Mie Keegan & Toshioka. 1957: 16. (as

//. incrmis) Coll. Asanunia. I 7 Jan

'54, (.idl . 27 Ml.

Vegetation Siiiniane Iwasaki el al.. 1964. las//. '62- iticrnm). Coll 1 suelile. .Sep Aug '6.V

Suglmolo. 19.17a:315. (as//. ttwrmts) Probably cited froni

Neumann { 19(16).

Cattle Shiinanc Sanhe KiIaoka& Morii, 1967a: 145-I52.

(I , M), Nov '62, (as ambi^w).

Capncurnis cnspus Neumann. 19(16:217, 1911:109. (Serowj (1 I) (as incrmis and ambigita. may be the same specimen).

Nutlall & Warburlon. 19 I 5: 367-36H. (as itwrmis). Cited Neumann's rect)rd.

Horsi ruki>ka Nuttall& Warburlon. 1 9 1 5: .t67-.!6X. (su) (= (as incrmis var. aponomvidcs)

I ukuoka) (1 I ). Berl. Mus. 17.1, also examined b> Hoogstraal. 1969.

I iikiiuka Keegan & loshioka. 1957: 16. (as incrmis var aponomindc\). (lied Irom Nutlall & Warburlon. 1915.

lloogslraal. I 96 1 :3| 7-31 8 (as incrmt\ aponttmoidcs)', 1962:195-

1 99 (as jpunoinoidcs). Cited from

Nutlall cV Warburlon. 1915.

Cow Shimane Sanbe lloogslraal, 1969:21 1-221 (as kitookai), Holotype, allotype, and paratypes.

Ccrxns nippon Mie Hoogstraal. 1969:21 1-221. Coll. (Sika Deer) Yamagutl, 22 Mar "67, (9 M, 19 11. Paratypes.

Horse and Hoogstraal. 1969:21 1-221. (2 II. Caprivonns crispus Identitied by Neumann ds amht^ia.

(Serowl Hrit. Mus. #1367. #139. (1 I ). Identitied by N. & W\ as incrmis vdT. apononioidcs. Berl Mus. 173. tlacmaphysalis Horse Kanunokuni =-55-J-()()S. Coll. losliioka* Aki-

longicornis lliyjiua yama. 17 Aug '55. iJ I').

#55-J-()32, Coll. Toshioka & Aki- !') yama, 17 Aug '55, ( 1 1.4

lomioka #55-J-(Hii. (nil Toshioka \ Aki- yama. 2 Aug '55. IV /•').

Kaininokuni. = 55-.l-ll49.( oil Tosliioka *: Aki-

Hi\ .iin.i yama. 17 Aug '55. 1/ /•')

Nagasaki lua/ato. Isushima :;67-J-(ll 211, Coll. liptonii Mi/u-

sawa.6 Mav '67,(52 I , 19 M, 9N,.M/. -/.V).

#67-J-()123.Coll. hploiiA Mi/u-

sawa, b May '67.(9 I . 3 M, X N.

/ F. I ^n.

Hokkaido Misaki. Abashiri #67-J-l)189. Coll. loshioka & Sasa-

gawa. 13 Jul '67, (15 I , 29 N,

.V /").

Korea I udii ri. Cbeju Do =67-K-(IIKil.( oil. Munsu.5 May '67, (.ff. 2N). Biological Series, Vol. 15, No. 1 Ticks 201 flacfnaphysalis Horse Korcu *ft7-K-()0()y. CoU. Munsu. 5 May longicumis •67, (.5 K h m.

Cattle Japan Hirosliima A?um;iy;iniij #55-J-()09. Coll. Toshioka & Aki- yuma. 19 .Sup "55. (/ Af).

#55-J-OI I. Coll. loshioka & Aki- yama, 19 Sep '55. (ft /•, .i M).

Hokkaido Migasliiniokoto, #.S5-J-(12.';.Coll. losliioka* Aki- .s.s,( Abashiri yania, 1.1 Aug 1 3 I , 9 N. 2J F).

Sliinlokli. iukaLhi #67-J-0193, Coll. losliioka&Sasa- pawa. 15 Jul '67. (4 1.2 N..

Nagasaki Oshinia-gun. Iki »67..|.|)II).S. ( ui: I iptun & Mi/u-

'67. _^ (K.) sawa. 4 May (6 1 , I').

Hatsuyama. Ikl # 67-J-(ll06.Coll. ripton& Mw.u-

(Iv) sawa. 4 May '67.(22 1 , 5 M.

42 N, / /'. ft A/. ft.V).

#67-J-010X, Coll. Tiplon ii Mi?u- sawa. 4 May '67,(6 !, 5S N.4 F. UN).

#67-J-() ill. Coll. Tipton & Mizu- sawa,4May '67,(1 1,7 N, J F,

/ M. UN').

#67-J-01l2.Coll, Tipton & Mizu- '67, sawa, 4 May I 3 1 . 27 N, 2 F. 2 M. 21 IV).

#67-J-()l 1.1. Coll. Tipton* Mizu-

sawa, 4 May '67,(4 1 ,4 M. U N, 23F.4UM. 7N).

Takcnotsuji. Iki #67-J-01 15, Coll. Tipton &. Mizu- (Is.) sawa. 4 May '67, (2 !, 4 M, 7 N, 2F.3M.8N).

^7-J-Ol 16, Coll. Tipton & Mizu-

sawa, 4 May '67,(1 N,i/-', / M.

I IV).

Gonoura. Iki (Is.) #67-J-014.1. Coll. Tipton & Mizu-

sawa. 2 May '67, (2 I . / Ft.

Iwazato, Tsushima #67-J-01 19. Coll. Tipton & Mizu-

sawa. 6 May '67.(2 1 . 1 1 N, / F. 211 m.

Kj^oshinij kliinan #67-J-lll27,Coll. lipton & Mi/u- sawa. 1(1 May '67. (/ Fl.

#67-J-ll 1.1(1. Coll. Iipton & Mi/u- '67. sawa. Id May ([ \ . 2 F).

#67-J-0 1.13, Coll. Tipton & Mizu- sawa. 10 May '67.(3 V.Jh').

#67.J.(1 136. Coll. liplon & Mi/u- sawa. 1(1 May '67. (J Fl,

Sakura Jinia »67-J-(ll39. Coll. Mizusawa, 14 May '67.1/ Fl.

Shinianc Sanbc #67-J-()l52,Coll, Holuhcc & Sugi-

yania. 15 May •67.(80 1 . 3 M,

28 N, / /!.

a66.J-(l()23.C oil iloluhii & loshioka, 13 Jul '66.1/ Fl.

#6(.-J-(l(l25.Coll. Ilolulx-i & loshioka. 13 Jul '66. (/ F"l

-66-J-0020. Coll. llolubccS loshioka. 6 ()Lt '66, 12 M. 2 AT.

#68-J-ll327. Coll. Yaiiiaguti & Hk-tha. 4 Jun '68.(2 1.4 M.65 N). 202 HHt(:nAM Doling Uninersity Science Bulletin

Uai'ttuiphysaUs Cattle Japan ShiiiianL' Sanlje #6«-J-0.t29, Coll. Yaniagutl & longicornis Kleeha, 4 Jun '(

lU'ppu. ok I (Is.) #66-J-l)026,Coli. Yaniagiiti. 12

Sep '66. (X I . .1 M. J2 N. 1110 L, J /•. .' ^f).

^6(i-J-()02l,(oll. Vainaglitl. 12 Sep 'M.il /••).

San be =hS-J-ll.V1(). Coll ^'ani.iguli&

HIeeha. 4 Jun 'hS. ( 15 I . 2 M.

1 Nl.

Ki'nuHlii Pasture. =h6-J-(lll2'),( oil, Yaniagutl. .1 Sep Takayaina-niura. '6ft. (/ F). Agatsuma-gun

=?fth-J-l)ll22. ( oil Yaniagutl, 3 Sep /•). ftft. ( /

Iwatc #67-J-OI4h, Coll Salto. 2h Apr

•62.(7()N. / /-'I.

Sotoyania = h7-J-l)2l I. Coll Yalllagull. Ill Aug '(i7, (17 1).

ff67-J-(l212. Coll "laniagutl. 10

'67. (>) Aug I . I Nl

#67-J-y2l.?.Coll Yaniagutl. 10

Aug"67.l« I . 2 Nl

Ua> asaka =f.7-J-(l214. (oil 'laniagutl. 1(1

Aug 'h7. (2 1 . 1 N. 1 Ll

lokyo H.iLhijn Koshnna 5h7-J-(l2ll7. (oil taniaguti. ]^

Jul'h7.(2 I . / /-'I

Aogoashinia Sh7-J-ll2(l>(. (oil lakallashl. 2ft '67. /) Jul ( Id I . /

Korea Ih.i-Ri.Cheiii Do Sft7-K-l)()l)2.Coll Yi Muiisu. .S Mav

ft7, |I4 N. / M. I .VI.

Ha-Gl, ( heju Do ?)!b7-K-ll003,Coll. Driggers. 5 May •hl.iJh: J At. /.V).

Naedo-Ki. ( lie|N Do ^7-K 0004. Coll. Dnggers. .S May

'h7. It, M).

#ft7-K-()00.'i. Coll Driggers. 5 May •67,(2 1 . I N.-4F\.

Jf ft7.K-0()06, Coll. Driggers. 5 May ft7.i.11-, 2 M.4K I M).

Ilalliin. Cheju Do #ft7-K-0007, Coll. Parsons. .S May ft7.(2 i: I M. /.V).

Bay-uni. Clieiu Do «h7-K-00()S. Coll Parsons. .S May

.S 67. (X I . N. / /. 2.VI

Song-C;a, Cheju Do •!;ft7-K-00l 0. ( oil Driggers, 5 May

•ft7,(.'; 1-. .1 M, 7 N, 7 h\ I M. :.V].

^67.K-I)0I l.( oil YiMunsu.5 May

'67.(2 I . 4 N. / K J m.

-67-K-1I012. (oil. Parsons. .S May

'67.(1 I-. 1 N.-//-. / .1/1

-67-K-(ini3, Coll. Yi Munsu, 5 Mav

'67,(4 P. 4N, //', I M. .<.V).

Aewol, Cheju Do #67-K-OOI4,Coll. Driggers. .s Mav

'67,(2 I-, 1 M, JF).

*67K-OOI5. Coll, Parsons, 5 May

1-, '67,(1 I M, 9 N, :f. J .W, BioLOGicAi, Series, Vol. 15, No. 1 Ticks 203

HacmapllvsaUs C 'at t Ic Nabum-Ni. Cheju Do #67-K-lMll6.Coll ViMunsu.S May

'67,(1 1-. I N. .'/•, J M. I ,V).

Kumik. Cheju Do #67-K-0()2(), Coll, Parsons, (. Mav '67,(4 I. 1 N,.;/-'l

Kumsong-Ni. Chejii =67-K-()021, Coll, Parsons, (, May

Do '67,(1 \. I M)

Sogwan. Cheju Dt) ir67-K-()(i:6, Coll Parsons, f, May '67. Cl 1 . : N. 2 h. .',1/1,

Kwason. C'heju Do #67-K-()()2X. Coll, Driggers. 6 May

'67.(1 M. 2 N. 2 1,.,V/--, / A7, 2 .V),

Ccrvits nippon J J pa II I othigi Kiiuigawa rf56-J-()6I.Coll, losliiokaS Akl- (Sika Deer) '56, yama. 25 1 eb (/ \U

Dog Hokkaido Misaki, Abasliin #67-J-(ll')II.Coll losliiokaii ,Sasa-

gawa, \2, Jul '67. ( 14 I . 5 N.JF),

Kozawa. Shiribeshi #67-J-(12()l.Coll. Toshioka& Sasa- gawa. IK Jul '67, (35 1-. 33 N, Ih F).

#67-J-0203, Coll, Toshioka & Sasa-

gawa. l(i Jur67,(/F),

#66-J-(>24(). Coll, Yamaguti. 3 Sep '66, (/ F. -',V|

lokyo Haehlju (is.) #67-J-()208,Coll. LItsumi. 13 Jul /•). '67,(2 1 , /

Donic'stiL- sheep Korea Ishidor ) arm. #67-K-0023, Coll, Driggers. 6 May Cheju Do '67,(1 N.5M).

#67-K-()()24. Coll, Driggers & Yi '67, Munsu, 6 May ( / K / A/1,

Vegetation Japan Hokkaido Misaki. Abashin #67-J-() 191, Coll, Toshioka & Sasa-

gawa, 13 Jul '67, (3 1 . 4 N. / F).

Korea Ch'onsong-Ni Hoogstraal et al,. 1 MhKl 2114- 1 2(16, Coll, Traub. 16 Jun '52. (1 11. RML #33(1'):,

Cattle, horse, slieep Cheju Do and Moogs'raal et al,. 196S: 12(14-1206. goat, dog Koje Do Coll, Parsons and/or Driggers and \i Munsu. Other data, see

Hoogstraal el al,. l')6H

Japan Hoogstraal et al.. 14(>S: 12(14-12116, Lcetotype temale, paraleetotype

male & synlypes, (6 1 , 2 M) N, 1425 in Neumann eolleetion,

Dog, eow K>ago i. Haragi Hoogstraal et al,, 1968: 12(14- 1206

(3 1, 2 M) marked "TYPi;", ( I 11 e\ iou. Kyago. (lb) e\ dog. Haragi. in Merlin Miiseuiu

Shizuoka Izu Hoogstraal el al,. 196K 12114-1206.

II 1 I lint, Mus, #1 I. 12, 13. 14.

1 2(16, CatllL- Kvushu Hoogstraal et al,. I 96S: 204-1 '33. Coll, Sugimoto. 14 Jul (4 1 .

1 Ml RML #151X5,

Horse Miva/aki Takanabe Hoogs(raal et al.. 196S 1 204- 1 206.

(9 I . 1 Ml N, 2409. Nuttall

( ollestion in Unt, Mus,. same speeimens m N, & VV,. 1915,

Tottori Daisen Hoogstraal el al,. 196X: 1204-I2()6. (X M N, 290X. Nuttall Colleetion in Brit, Mus,. same speeimens in N.& W,. 1915,

Cattle. hor>e. dog. Hokkaido Hoogstraal el al,. 196X, badger, deer At)mori Iwate Gumma Nagano 204 Bhigham Young University Science Bulletin

HaemaphysaUs Cattle, horse, dog. Japan Tokyo Hoogstraalct al.. 1968. longicornis badger, deer Naru Hiroshima Shimane Nagasaki Kagoshima

Cattle, horse, dog various provinces

Dog Korea

Man Japan Kyoto

I.cpus hrathyunis Nagano brachvunis (Hare)

Cattle

Kagoshima

Okayama

Cattle and horse Hokkaido

Horse

Vegetation

Tokyo

Cattle, horse, sheep, Japan dog, goat Biological Series, Vol. 15, No. 1 Ticks 205

Haetnaphysails t aitk\ horse, dog. Japan Shiinane Itagaki. Noda & YainagULhi, 1944. longicornis sheep, goat Oila 1955. (as//, hispinosa). Korea Miya/aki Kagoshima Hamgyong- pukto

Korea Hamgyong- naindo, Kyonggi-do. Kyongsang- puklo, Kyongsang- nariido. Cholla- nanido

Cattle, ht»rse. human, Japan Honshu. K\ iisfui Hi Sugimoio. 1936e:580. (as//. wild rabhil & Hokkaido hispinosa). Korea

C attic & horsi Japan lukuoka, Sugimoto. I937b:605. (as//. Kumamoto hispinosa) (F. M, N. L). Jul-Oct Miyazaki '33 and Jul-Nov '36. Kagoshima

Korea all over the peninsula

Lepus hrachyums Japan Niigata Kakudayama. angiisttdens (Hare), Meles Sasagamme Pasture

meles anakuma ( Badger). Si. Ohsadorynkan \'yctereu!es procvonoides Pasture viverrimts (Raeeoon Dog), cattle & vegetation

Cattle, horse. Ai. hare Japan

Cattle, horse, goat

Cattle, horse

Cattle, horse, hare. Sox. &. vegetation

Horse 206 Rrigiiam Young University Science Bulletin

Hacmaphysalis Horse Japjn NulIalUSc Warlnirlun. 1415:433.

longicornis (as//, htspinosa) I A. I.) N. 19()«.

' 2 Aug 1 2. tnund with IxoJcs

ricmus (- ' /. nipp(mcrisis). also examined hy Hoog\traal.

Miya/aki Takanabf Nuttall& Warburton. 1915:433.

(a-.//, hispmosa) (I ) N. 1909, 5 Jun 12 Oct '1 2. aKo cNannned by Hoogstraal.

Cuttle Tokyo Nuttall& Wurburlon. 1915:433.

(av//. hispmosa) (1 1 ) N. 29 11 a.

1 6 Oct '11, tbund with Hoophilus \p,. submitted by Dr. Miyajima.

Hacmaphysalis Sus scrofa leuiomyslax Mic Coll, Yamagiiti. N Mar 'hi. (ISM). mcgaspinosa (Wild pig)

Cen\(s nippon Coll. Yamaguli. Mar 7i7. (4 1 .

(Sikj Dc'cr) 1 9 M ).

Shiga Coll 'laiiiaguti. Mar 'Ul . ( 1 } M).

Miya/aki Kitagawa-rmiru & Coll. Vajinia. 15 Mar '61.(2 i). Aya-machi, Higashi- rnorokata-gun

'68. K a ruga w a Miyagasc Coll. Kilaoka. 27 Nov (31 I . 15 M.X4 N.8 L).

Coll. Kitaoka. 5 Jan-23 I eb '69. (Numerous adults)

Capricornis crispus (Serow)

Vegetation Nanasawa. Alsugi- machi, Naka-gun

Mmeyama. Kuribara. Isehara-machi, Naka- giin

Hacmaphysalis Pcnlalam s furncs-ii Ryukyu pentalagi (Ryukyu Rablntl

Haemaphy salts wellingloni I Biological Series, Vol. 15, No. 1 Ticks

Ixodes acu litarsu i M a n J a [la n

Capricorn is crispus (Serow)

Serow. cattle & dog

Ixodes an^istus Clcthrionomys rufocanus bcdfordiae (Rcd-backctl mouse!

Vegetation

Clvthnonornys rut this (Red-backed mouse)

Apodemus speciosus aimi

(I leld mouse)

Clethnonomys rufocanus

bedfordiae ( Red-backed mouse) & Apodemus speciosus ainu (Meld mouse)

,, Clethnonomys mfceanus bedfordiae (Red-backed mouse)

Clcthrionomys rufocanus hedjordiae (Red-backed mouse), Apodemus speciosus ainu (Field mow^c). Apo- demus argenteus hokkaidi (Field mouse)

Ixodes granulafus Rati us raitus (Rat)

Suncus murinus nukiuanus (Shrew)

Rattus norvcgieus (Norway Rat)

Rattus sp. (Rat)

Under log 208 Bhigham Young Univehsitv Sc:ience Bulletin

Ixodes gramilalus Railiis rallus i Rat I Ryukyu Okinawa Kcegan & Tinhioku. 1957; 19. Coll. Islumls Posekany, 29 Apr '45. RMI, #27512 nil.

Kcugan& Toshioka. 1 957: 19. Coll.

Povekany. 5 May '45, (2 1 -RML

^27514, ,1 H-RML #275 13).

Kccgan& I oshioka. 1957: 19. Coll.

Posekany, 16 May MS. RML #275 I S

(2 1,2 M, I Nl.

Kccgan &. 1 oshioka. 1957: 19. Coll. Hardcasllc, 10 May 45. RML

s:275l()(4 1 ).

>'jinamiwj. Okinawa KcL-gan& Toshioka. 1957: 19. Coll. Hardiastlc. 2.1 May '45. RML #27519(3 V).

Nago, Okinawa Kecgan & Toshioka. 1957:19. Coll. Hardcasllc. May '45. RML #27521

(I I ).

Naha. Okinawa Kawashiina. I9(,.l: 1(1.1. Coll.

Kawashinia. 25 Jun '62. ( I I ).

Japan Tokyo llachijo (Is.) .^sanunia& Sckikawa, 1952:107-1 16, 1953:99-1 12. Coll, Asanurna,

Dcv "51 & Mav '52, (A. N. 1.1.

Miyaki (Is.) Asanunia& Sekikawa, 195 2:107-1 16, 1953:99-112, Coll. Asanuma, Mar '52. (N.LI.

Rattus noryegicus llachijo Koshinia Asanuma & .Sekikawa. 1952: 107-1 16. (Norway Rat) 1953:99-1 12. Coll. Asanuma. Sep '50 & Aug '51, (N, L).

Asanuma* Sckikawa, 1952:107-116. 1953:99-1 12. Coll. Asanuma. May '50 & May '52. (A).

lachljo (Is.) Asanuma & Sckikawa. 1952: 107-1 16. 195 3:99-112. Coll. Asanuma,

Mar '50. Aug'51.& May '5 2. (A. N. L).

Miyaki (Is.) Asanuma & Sckikawa. 1952:107-1 16. 1953:99-1 12. Coll. Asanuma. Mar "52. (N. L).

Kochi OkmoShima Asanuma & Sckikawa. 1 952: 107- 1 1 6. 1953:99-112. Coll. Asanuma. Aug '52. (Al.

ApoJcniu^ \pi'ti(tsus Tokyo Mi>akc(ls.l Asanuma & Sckikawa. 1952: 107-1 16.

1 1 icid Mouse) 1953:99-112. Coll. Asanuma. Mar '52. (N. L).

Chihara Asanuma & Sckikawa. 1952:107-1 16. 1953:99-1 12. Coll. Asanuma. Jan '53. (N).

Kanagawa Kurihama Asanuma* Sckikawa. 1952:107-116. 1953:99-1 12, (oil, Asanuma. Jan'53.(Ll.

Dog& car Tokyo 1/u-shichito Asanumai. Sckikawa. 1953:99-1 12. Coll. Asanuma. (2 A).

Siincus muriiuts (Shrew) R>uk,\u Nago. Okinawa Kecgan it loshioka, 1957: 19, Islands Collected 26 May '45.(1 II.

I'urdus cchcru>pus Japan lokyo Miyakc (Is.) Asanuma & Kosaka, 1955: 195. felat'nopiis Collected (II. (Seven Islands Ihrushl

Apodcmus ayronus ssp. Paedun-ni. Panstmg. Arthur. 1957:6N3-6S6. Coll. (lield mouse) Chindong-ni 37th Med. Co.. Jun & Jul '54.

(lOM. 30 1 ).

Ixodes lividus Di'lichofl urhica dasypus J:ipan Shimanc lliiioinisaki -66-J-(l027. Coll. Toshioka. Hctch- (Japanese House Martini Icy & Holubec. 9-15 Jul '66. (.5 /•"). Biological Series, Vol. 15, No. 1 TlCK.S 209

Ixodes lividus Dclkhoti iirhua Jasypus Japan Guriima Minakanii (Japjncsc Mouse Mjrtin)

Riparia riparia ijimac Hokkaido Hamamokoto. (Fastern Sand Martin) Abashiri-shi

Delichon urbica Jasypus Nagano (Japanese House Martin)

Ixodes Vegetation monospinosus

Man

Ixodes nipponensis Cattle

Cenus nippon Mic (Sika Deer)

Dog

Lcpus hrachyurm

aripislidens i Hare \

Vegetation

Dog Kochi

I.epus hrachyums Niigatu

angitsiidens i Hare)

( altle. horse, dog, badger, hare, weasel, man. small rodents & birds. 210 Rmu;ham ^'oung University Science Bulletin

Ixodes nipponensis Clethriononiys mfocanus J J pan Hokkaido [shikan ^: )lulaka Ono. 1966:62-68. Coll. Ono. Oct

hvdfordiac ) Rcd-buLkcd '55 - Aug '62. (N. L). as Ixodes nicnisc). C. ntiilus mikado sp. NA & I-A. (Rcd-bucked mouse), Apodcmus spcciosus amu

{McUi mouse). ,-1, ar^cn lens (licid mouse).

Dog Miyjgi

Rodenis &. shrews

Ixodes ovatus Cut lie s

Biological Series, V'ol. 15, No. 1 Ticks 211

Ixodes ovatus Vcticlalioii J^ipan Hokkaido Tsukisuppu. #55-J-()47. (oil. Toshioka & Aki- Ishikari yuma, 5 Jun '55, (/ F),

Apoiletnus spcci(}sus aimt Teinc. Islnk.m (\)ll Orui. IN Oct "55. (/ A). (licld inuLisc)

Clethriononiys mfociuius Coll. Olio. l^J Apr '5(1. (/ ,V). ht'dfordiav (RL-d-backfd nunisc)

Caltli.', hor^c & human Hakodate, Sapporo, Ogura & lakada, 1927;201-2()3

Muroran, Atsunai (as /. jrequem) {V, M).

Horse, cattle, dog, hare. A kit a. Kishida. I93()a; 1-5. as /. & human Shi/uoka, frcquens. ovatus. japoriensis. & Kyoto, carinatus. Record oi Jrcqucns Saga, is cited from Ogura & lakada Tokyo. (1927). Kanagawu, Hokkaido

Cattle & luirse Niigata, Nakaniura& Vajimu, iy37:145-MK. Akitu. as japonerisis & frequens. Shimane, Hyogo. (lumma, rukushima. Hokkaido

Aomori, Yajinij. 1442:507-504. ds frequens. Iwate

Aomori. Itagaki, Noda & Yamaguchi, IM44, Iwate, 1959, ds frequens. rukushima, Miyagi. Nagano. Hokkaido

Man. cattle, dog. cat, Niigata Niigata Pret. in Sailo, 1959b: 199-203. 1964:60;

Mustela sibirica itatsi Honshu Saitoet al., 1965:145-146. (as

(weasel). Xvclcrculcs /, japonensis) Coll. Saito. Jul procyonoidcs viicrrinu & '55- 17 May "65, (A. N, L). (Raccoon Dogj. Moycra wogitra \vogi4ra (Mole), Sado (Is.) I.i-pus hrachyunm lyoni Immature forms on small rodents. (Hare), Raltus noncgi- cwA(Nor\va> Rul), f.i'pus hrachyunis anfiustidens (Hare), Mtcrotus monte- belli (Korean Meadow Kanagawa Mt. Ketsu, Vole), M. monicht'lli Iseharacho

brcvicorpus ( Meadow Vole). Apodcmus spcci-

osus sadoensis (i ield mouse). Crocidura dsinezumi (Shrew), & vegetation

Rodents & shrews Hokkaido Supporo-sln Ono. 1962b: 155. (as/, sp. 6) Coll. Ono. (L).

Clclhnonomys mfocanus Oshima, Ishikari. Ono. 1966:62-68. (as/, sp. 6 & bedfordiae (Red-backed Hidaka, Abashiri, 52) Coll. Ono. Jun '55 - Oct *62. mouse), C. mtilus Tokachi. Rumoi (N.L). mikado (Red-backed mouse), Apodcmus argcnteus (1 ield mouse). A. speciosus amu (Field mouse).

Clethrionomys rufovanu s leme Asanuma, 1955: 1 240-1242. (as /. bedfordiae (Red-backed sp. 52) (N.). mouse).

Apodcmus spceiosus Aomori, Asanuma & Sekikawa. 1952:107-1 16. (I-ield mouse), Yamanashi, 1953:99. (as/, sp. 6 & 52)

A. argenleus ( lield Chiba. (N.L). mouse). Hothenomys Kanugawa. smilhi (Vole). Mus Toyama. museulus (House i'ukui, mouse), Mierotus Shizuoka. montehelli (Korean Kyoto, Meadow Vole) Gita. 212 Bhicham Young Univehsity Science Bulletin

Ixodes ovatus Dop & hare Japan Chiba, Asanuma. 1956:92-93. (as/.

Akila. iap'iuviiMs) {A I, lukushima, Miyagi,

Nilgata Kcctian & Toshioka. I957;2(l. (as

/- japoyjcnsis) Coll. Saito, 27 Jul 55.(2 II.

C'altk Okayama Kecgan & loshioka. 1957:2(1. (as

/. japunftisn) Coll. Vajnna. 1 ^ Jul

•55. ( 1 1 ).

Ktcgan & Toshioka, 1957:2(1. (as

/. japom'iisis) Coll. Yajinia, 10 Jul

'36. (111. (labflli-d d-. frcQiu'ils).

Dog Niigata Gini'undaira Kccgan & Toshioka. 1957:20. (as

/, japonensis) Coll. Sailo, 31 Jul '56.(1 Ml.

Ratlits nurvc^lcus KcLgan & losliioka. 1957:20. (as (Norway Rat) /. japoni-nsis) Coll. .Saito, 26 Jul

'55,(1 1 ).

/ cjuts hrui hytinis Nagano K.iniiko^hi kccgan & loshioka. 1957:20. (as

/. ) llironiatsu. brat hvttruK ( Marc) /aponcrisis Coti. 4

IX-c '52,(1 1 , ION),

Kochi Kccgan & Toshioka. 1957:20. (as

/. lapont-nsis) Coll. Hironiatsu.

21 1 cb '56.(5 I-, 1 Ml.

Man Tokyo Kccgan & Toshioka. 1957:26. (as

/. japoiu'iisis) Coll. Kishida, (III. (Hoiotypc ol cannatus Kisliida. 19301.

Horse, hare, dog Akita, Nciniiann. 1S99:1 16-1 IX. Coll, Saga lagULlli (Akital & Yaniaguehl

(Sagal. (4 1 I. Type scries

contained 2 I of r/c7>(MS (=? pcrsitlcatus or mppotwnsis).

Horse, dog Saga Neumann, 1904:452-453. 191 1 U&

Diinitz. 1905:132, (2 1 ol 4 1

above I.

Nuttall& Warburton. 191i:15S. (2 II. As ricinus var. ovatus.

same specimens as Neumann I 19041

Tokyo ? near Tokvo .Neumann. 1904:45X. 191 1:22.

Nuttall& Warburton, 191 1 :20>l-209. (as/, japortf lists) Coll, Harmand,

1901,( 1 1 I, type specimen, (Pans Museiiml.

Horse, dog Saga Morel. 1963:925-928. (2 I I. same

speciniens as Neumann ( 19041. Morel synonymi/ed japonensis under ovattts

Hokkaido Shikotsu lake Schul/e, 1930:294-303. (as/ Iburi frequens) Coll, S. Schocde. 16 Jul '06,(1 Ml.

Ixodes persulcattis Cattle Shintoku, Tokachi #67-J-0197.Coll. laniguchi, 17 May '(.l.ifi F. IJM. I A).

,i Kuttan, lokadii Sr67-J-0199. Coll. laniguchi. I May '67,(i/-'l.

Ko/awa. Shiribcshi #67-J-0202, Coll, Toshioka & Sasapawa. IS Jul '67, (/ /•'!.

*67-J-(l242.Coll, Yaniaguli.6 Jur67.(/ F).

Hor\c Hokkaido Shintoku. lokathi =67-J-OI95, Coll, laniguchi. 10 May '67.1/ /•).

Shi/unai. Hidaka #67-J-0200. Coll loshioka &

Sasagawa. 16 Jul '67. iS /-"I. Biological Series, Vol. 15, No. 1 Ticks 213

Ixodes fXTsulcatus Horse Japan Hokkaido Sliiraniika. Ku'ihiro #55-J-05 I . Co!!. Toshioka &

/•"). Akiyamu. 14 Aug '55. ( J

Vrsin arctos yt'sot'nsis Shintoku, Tokachi #670-0196. Coll. Taniguclu. 19 (Bear) Jun "67. (i/-; 2 m.

Dog Kozawa, Shiribcshi ff67-J-02f)4. Coll. Toshioka &

Sasagawa. IS Jul '67, (/ f-\ I M).

#67-J-0:44, Coll. Yaniaguli, 25

Apr "67, (/ h').

Man Tokushimj Mt. 1 surugi Keegan& loshioka. 1957:21-22. Coll. Tanaka, 16 Jur56.(2 \\

1 N).

Horse I wale Keegan & Toshioka. 1957:21-22. Coll. Yajima. 27 Jun\l«.(l !,

1 M).

Dog Niigata Ginzandaira Keegan & loshioka. 1957:21-22. Coll.Saito. 25 Jur55.(2 1).

I.epus hrachvurns KoLhi Aki-shi Keegan & Toshioka. 1957:21-22. hrachvums (Hard Coll. Hiromatsu, \^ Nov '55-10 Mar "56. (I\ M).

Weasel Keegan & Toshioka. 1957:21-22.

Coll, Hiromatsu. (1 I ).

Dog, Lt'pus brachyums Niigata Niigata Pret. in Saito. 1969b. 199-20.?. Coll.

- angiislidens (Hare), Honshu Saito. Jul'55 I eb '59.

/.. braihyunts lyoni & (A.N.L), (Hare), eattlc. Sado (Is.) Nyctcrcutes procyonoides (Raeeoon Do'^), Micro tus man [c belli brcvicorpus Kanagawa Mt. Ketsu. Isehara- (Meadow Vole). Apodemus cho spccioaus sajocnsis (lield mouse). Rattus norrt'^f^s (Norway Rat), Croadura dainczumi (Shrew). Phasianus colchicus rubu stipes (Pheasant), i vegetation

Rodents &. shrews Hokkaido Sapporo-shi Ono. 1962b: 155. Coll. Ono, (L) (as Ixodes sp. 2).

Clcthrionomys ntfoeanus Oshima, Ishikari. Ono. 1966:62-6S. Coll, Ono. Jun bedjordiae (Red-backed Hidaka. Soraehi, '55 -Oet '62. (N. L). mouse), C nifihis Kaniikawa, Tokaehi, mikado (Red-batked mouse Kushiro. Nemuro, Apodemus speciosus Abashiri.

amu ( lield mouse). A. ar^enteus (I ield mouse).

Clethnonomys ntjoeamis Shintoku, Hiioo. Asanuma. 1955:1240-1242. (N. L). as sp. 2 54. bcdfordiae { Red-batked Nohoro, Sounkyo, Ixodes & mouse). Apodemus specio- Otoineppu, Toyotomi, sus aimt (Field mouse), Shiranuka. Peine. A. argenteus hokkaidi Nonaka.

il ield mouse). Mus

musculus yesonis ( House mouse).

Apodemus speciosus Hokkaido Asanuma & Sekikawa. 1952:111, Coll. airw (I ield mouse). Aomori Sugayu 1953:99 & 105. Asanuma. "53 A. argentues (Field Akitj Tazawa Ma> -Jun "53. (N. L), as sp. mouse), ,4. speciosus Saitamu Chiehibu Ixodes 2 & 54. (Field mouse), Yamanashi Yamanashi & Yamanaka holhenomys smilhi (Vole). (lunniij O/e Cicihnonomys rufoeamis (Red-backed mouse), Mus musculus yesonis (House mouse)

Hokkaido. Kitaoka& Saito. 1967:82. Niigata, Akita, Yamagatu. Nagano. Crumma. Gilu. 214 Bricham Young University Science Bulletin

Ixodes pcrsuhalus Japan Hokkaido Shikolsti lake. Schul/e. !y,H»: 294-303. Coll.

Iburi Schoede. 16 Jul '06. (2 I ).

Htirsc & hare Akita. Neumann. 1899:116. 1904:452; Saga NuttallA Warhurton. 1911:156. (as/, nanus) Nuttall & Warburton cited Neumann's record.

Varunis aminals Neumann. I'M 1 1 M 3. (as/. ncinus).

Horse Dai sen Nuttall & Warhurton. 1415:433.

(as /. ricinus) lound with //.

bispinosa {=11. longtcornis) . 1 Aug '12.

Oiltlc. horse &. nun Aornon. Ogura& Takada. 1927:201. (as

Hokkaido. /. rieinus) (I . M).

Human. i.attle. horse, Hokkaido. KIshida, 193()a:2.(as/ ricinus) hare, deer. dog. fox. Tokyo, roe-deer & leopard Nagano, (troni /uo '). marten, Kyoto. hedgehog, weasel. Hyogo, sheep, goal. L-'hime, Saga.

Korea N. Korea Kishida, 1916: 142. (as/. ruinusl

Horse ? Japan Miya/.dki Sugimoto, i937h:603-6()4. (7 I ). ds ricinus var. mivazakicnsis.

Horse, cattle, dog Kumamolo.

Korea

(altle. horse Jap.in

Cattle, horse, dog

Korea

Ixodes philipi hijjinus ieucomclas or Japan Occanadroma caslro

Ixodes sigTiatus l.ants erassirostris Aornon (HIack-tailed C^ull)

Phalacrocorax pela^- cus (Pelagic Shag) Biological Series, Vol. 15, No. 1 Ticks 213

Ixodes si^ialus Phalacrocorax pcla^- Japan Yc/o Nultall& Warburton. 191 l:261-2ft4. cus (Pelagic Shag) (= Hokkaido) As sifjnarns. L*\aininL'ii Neumann's specimens as well as sif^ialus sent them by Hirula. and tound them identical.

Zumpt. 1952: IH. Merely lisis the previous record.

/.arus crassirosrris Aomori Kabujima. Hachinohc- Asanuma. Okubo & lukiida, 1955; (Black-tailed Gul!) shi K5-K6. Cull, lukudu & Asanuma. May '51 - Ma> "56. (A.N. L).

Crevices in rocks near Kabujima. Hachinohc- Keegan & Toshioka. 1957:22. Coll. nests ot Lams crassi shi Toshioka & Akiyama. IS Aug '55. rostns (Black-tailed (70 I, 53 M, 3 N).4()6 MGL ^H.

Gull) Coll. Akiyama, 14 Oct '55. (96 I . 72 M. II N). 406 MGL #40.

PhalacroLorax pclagi- Hokkaido Asanuiiij Si I ukud.i. 1957 147-159. ctts (Pelagic Shag)

Pfialacrocorax capillalus Hokkaido. Asanuma, 1965a: 1 16. (Temminck's Cormorant) Kanagawa

Japanese Cormorant Hokkaido Kishida. I930a:2. Kishida did not reter to the scientific name of the host. May be citation from Neumann or Nuttall & Warburton.

Ixodes simplex PipistrcUus ahramus Okayama Coll. Yamaguti. 9 Mar '57,(1 F). (Bat)

Myotis mavrodaetylus Iwate Coil. Lndo. 26 Jul '67. (/ ;V). (Little Brown Bat)

Bat Nagano Kishida, 19.^()a:.l.

Shiga Keegan & Toshioka. 1957:23-24.

Coll. Asanuma, ( I I).

Myotis macradac(ylu\ near Kameaka Arthur. 195^: 1S9. 30 Sep "52,

(Little Brown Bat) (I I , 3 L), RML #30943.

Minioptcnts sehrei- Kanagawa Kamakura Arthur. 1956: 1H9. 24 Dec '45. hersi (Long-winged (I N). RML #22366. Bat)

Ixodes tanuki Nyctercuies proeyonoi- Niigata Asahi-nmra. Ivva- Saito, 1964:59-66. Coll. Saito, des viverrimts (Raccoon tune-gun 23 Nov '59. (I F). Dog)

Kurokuwa-mura, Saito. 1964:59-66. Coll. Saito.

Kitakambara-gun 22 0ct '59, (2 1-. I M).(l M:

Holotype, I liAllotypc).

Sasagami-mura. Saito, 1964:59-66. Coll. Saito, Kitakambara-gun 23 Nov '63.(1 V).

Ixodes turdus Turdus celaenopus Tokyo I/u-shichito #55-J-024. Coll. Asanuma, 13 eclaenopus (Seven Aug '55. (4 N, IF). Islands Thrush)

Motacilla cmerea easpi- Yamaguchi Tokuyama-^hi Kitaoka's C'ollection, 26 Nov '66. ca (Eastern Grey Wagtail) (3 l).

Turdus celaenopus Tokyo Hachijo (Is.) Nakatsiiji. 1942:291-295. Coll. celaenopus (Seven Tono. 25 Dec '40. (7 I ). Islands Thrush) Keegan & loshioka. 1957:24. Coll. Asanuma. 12 Dec '55.(1 I. 6N). 406 MGL # i4K.

Aogashima Keegan & Toshioka. 1957:24. Coll.

Asanuma. Nov '54.(1 I, 1 N).

Turdus c. celaenopus „ (Seven Islands Ihrush), Turdus ehrysolaus ehry- \olans (Japanese Brown Thrush), k'rnheriza ele^ans

elegans ( Yellow-throated Bunting). Emhertza eioides ciopsis (Japanese Meadow 216 Rkicham Young University Science Bulletin

Ixodes turJus Hunting). Efnbcriza spodo- Japan Shi/uoka cvphala pcrsonata (Jupancsc Hunting)

Ccttia diphonv cantons Asanunia & Seikjkawa, 1 45 2: 1 I I . as

Warbler). I (JapancsL- Hush Ixodes sp . , Asaniirna & Kosaka. Ccttia diphouc ijimac 1955:194: Asanunia & Nakagawj.

{Jjpjncsc Hush Warbler). 1955:549. .11 May '51. Jan "52. Bambusicola thoracica Ieb& May '5.1. Jan - Mar & Nov '54. thoracica (I . N. L). (Chinese Hjmboo Phea- sant), Coccothratistcs cocco- [firaustcs japonicus (Japanese Hawfineh). Hypsipctcs atnaurotts amaurotis (Hrown-eared Hulbul). Pyrrhula pyrrhiila jiriscivcNlris (Japanese HuIIIinLh). Pfiylloscopus ijimac (Ijima's willow warbler) Apodcmus spcciosus (I ield mouse)

IxoUcs uhac ,\'yctalus sp. Niigjij (Hat) Nagano

l.unda cirrhata Hokkaido

( Tufted puttin)

Ixodes Rhinolophus jcrrum iothigi vcspcrtilionis cquinum nippon (Morseslioe bat) liioLOGicAL Series, Vol. 15, No. 1 Ticks 217

IxoiJcs Rhinolophus fcmun- Japan lochigi Kccgan*. Tiisliioka. 1457:25. ColL icspcrlilionis cquinum loshioka & Akiyanu, 25 Icb '56 (Ho^sc^hut• hat) ( I N. 4 L). 4(16 M(iL #191.

l.t'pus h. hraihvums Nayano Kccgan & liKhiiika. 1457:25. Coll. (Hare) HironialMi. 2 Mar '28, (IN,.? L). This rciord seems to be doubtful.

Rhinolophus fvrnim- Myiagi (sic) Arthur, 1956: 1X4; lloogstraal, 1956: cquinum (= Miyagi) 569; Kcegan& loshioka, 1957:25. (Horsosiioi.' bat) Coll, Nieholson. I.? May '52, (I I), RML#.12112.

Rhipiccphalus Dog Ryukyu Okinawa #66-R-()(l05. Coll, Doucette & san^iinvus Islands .Sugiyania. 19 Oct '66, {4 F. / ^T).

(amp McrLV. #6K-K-II(IIK, ( oil Hubert, .10 Sep

Okinawa '68,(2 I ).

Japan Kiinagawa Yokohaina-shi «64-J-llll79, (oil. I lint, 7 Get

'64, (2S I , 75 Ml.

Ogaishi-inura, Aso- Sugimoto, 19.17b:6l()-6 12. Sep '.1.1, gun (.11).

Kunianioto-shi Sugiino((i. I9.17h 61(1-612, l8Jul •.^5,(.l 1,5 Ml,

Man Osaka-shi Keegan & loshioka, 1957:26-27, Coll, Holland, Sep '56,(1 1).

Numbers in p.irentheses represent specimens senl (o Mi (;ien \1 Kolils lor contirniation ol delermin.ilions 218 Bhicham Young University Science Bulletin

APPENDIX 3

List of Maps and Text Figures

Map HioLocicAL Skuien, 220 Brigham Voung University Science Bulletin

98. Illustration of Ixodes ovatus. larva 140 99. Illustration of Ixodes pcrsulcalus. female 143 100. Illustration of Ixodes persulcatus, male 144 101. Illustration of Ixodes persulcatus, nymph 145 102. Illustration of Ixodes persulcatus, larva 146 103. Illustration of Ixodes philipi. female 149 104. Illustration of Ixodes si^natus, female 151 105. Illustration of Ixodes signatus. male 152 106. Illustration of Ixodes signatus. nymph 153 107. Illustration of Ixodes signatus, larva 154 108. Illustration of Ixodes simplex simplex, female 156 109. Illustration of Ixodes simplex simplex, nymph 157

1 10. Illustration of Ixodes lanuki, female 159 111. Illustration of Ixodes tanuki, male 160

1 12. Illustration of Ixodes turdus, female 162 113. Illustration of Ixodes turdus. nymph 163

1 14. Illustration of Ixodes turdus, larva 164 115. Illustration of Ixodes vespertilionis, female 166 116. Illustration of Ixodes vespertilionis, male 167 117. Illustration of Ixodes vespertilionis. nymph 168 118. Illustration of Ixodes vespertilionis. larva 169 119. Illustration of Rhipicephalus sanguineus, female 171 120. Illustration of Rhipicephalus sanguineus, male 172 121. Illustration of Rhipicephalus sanguineus, nymph 173 122. Illustration of Rhipicephalus sanguineus, larva 174 321

Biological Sekies, Vol. 15, No. 1 Ticks 221

APPENDIX 4

Information on Specimens Illustrated

Figure Species Remarks

- 5 Argas japonicus Adult from Coll. #66-J-0037 From nest of Hirundo daurica japonica (Japanese Striated Swallow), Niimi, Okayama Pref., Japan, 21 September 1966, Hatoyama & Mizusawa. Holotype female & allotype male, a-d, female; e-g, male, a, whole body (dorsal & ventral view); b, e, capitulum (ventral view); c, f, genital area; d, g, tarsi MV (lateral view). Adapted from Yamaguti et al., (1968).

Argas japonicus Larva & nymph from Coll. #66-J-0037 - Same as above, a, b, larva; c, d, nymph, a, whole body (dorsal & ventral view); b, Haller's organ; c, capitulum; d, tarsi I-IV (lateral view).

Argas vespertilionis Female from Coll. #67-J-0215 - From Vespertilio superans (frosted bat), Shojo, Fukushima Pref., Japan, 11 August 1967, Yamaguti. a-g, female: h. genital area of male, a, whole body (dorsal & ventral view); b, periphery (lateral view); c, posterior quadrant; d, tarsi I-IV (lateral view); e, spiracular plate; f, capitulum (ventral view); g, anal valve.

Argas vespertilionis Larva - From Pipistrellus abramus (Bat), Fukuoka-shi, Japan, 25 June 1967, Wada's collection.

Ornithodoros capensis Adult from Coll. #66-1-0054 - Under rocks near nest of Larus crassirostris (Black- Tailed Gull), Kyojima, Shimane Pref., Japan, 9 July 1966, Toshioka & Betchley. a-d, female: e-g, male, a, whole body (dorsal & ventral view); b. e, capitulum (ventral

view); c, f, genital area; d, g. tarsi 1-IV (lateral view).

Coll. - From Calonectris leucomelas (Streaked 10 Ornithodoros capensis Larva & nymph from #66-J-0054 Shearwater), Ohakajima, Shimane Pref., Japan, 13 September 1966, Yamaguti's collection (larva); #66-1-0054 (nymph), a, b, larva: c, d, nymph, a, whole body (dorsal & ventral view); b, Haller's organ; c, capitulum (ventral view); d, tarsi I-IV (lateral view).

- flavomarginata .4 geoemydae Female from 406 MGL CoU. #57 From Cyclemys flavomarginata 1 mblyomma (Turtle), Futanaka, Ishigaki (Is.), Ryukyu Islands, 6 November 1955, Keegan & Toshioka. Reproduced from Keegan & Toshioka (1957), pi. 4.

geoemydae Male from 406 MGL CoU. #57 - Same as above. Reproduced from Keegan & 1 A mblyomma Toshioka (1957), pi. 5.

Coll.#68-R-0015 - From Geoemyda spengleri japonica (Turtle), Yona, 1 A m hlyom ma geoemydae Nymph from Okinawa, Ryukyu Islands, .April 1968, Mizusawa.

14 A mblyomma geoemydae Larva from Coll.#68-R-O015 - Same as above.

- Snake. Naha, Okinawa, Ryukyu 15 Amhlyomma nitidum Female from RML Coll. #35991 From Sea Islands.

- near Inoda, Ishigaki (Is.), 16 A mblyomma testudinanum Female from Coll. #55-R-017 From Wild pig, mountains Ryukyu Islands, 4 November 1955, Keegan & Toshioka. Reproduced from Keegan & Toshioka (1957), pi. 6.

- 1955. Reproduced 17 Amblyomma tcstudinarium Male from 406 MGL CoU. #82 Same as above. 2 November from Keegan & Toshioka (1957). pL 7.

- scrofa leucomystax (Wild pig), Yuwan, 18 Amblyomma testudinarium Nymph from Coll. »68-J-0318 From Sus Amami Oshima, Kagoshima Pref., Japan, 11 April 1968, Mizusawa.

- vegetation, Yuwan, Amami Oshimu, Kagoshima 19 Amblyomma testudinarium Larva from CoU.#68-J-0317 From Pref., Japan. 10 April 1968. Mizusawa.

Coll. #73 - From cow, Hateruma (Is.), Ryukyu Islands. 20 Boophilus microplus Female from 406 MGL Reproduced from Keegan & Toshioka (1957), pi. 8.

pi. 9. microplus Male - Same as above. Reproduced from Keegan & Toshioka ( 1957). 2 1 Boophilus

CoU. #67-J-0261 - From cow, Yuwan, Amami Oshima. Kagoshima 22 Boophilus microplus Nymph from Pref., Japan. 3 October 1967, Kimura & Mizusawa. Bkic.ham Vounc University Sc ience Bulletin

23 lioophiliis microplus Larva from Coll. #67-R-0011 - iTom cow, Hirai, Ishigaki (Is.), Ryukyu Islands, 8 February 1967, Tipton.

24 Dermaccntor sp. Female - From horse, Unggi. North Korea, June 1942, Asanuma's collection. Reproduced from Keegan & Toshioka (1957), pi. 10.

25 Dermaccnlor sp. Male - Same as above. Reproduced from Keegan & Toshioka ( 1957), pi. 11.

26 Dermaccnlor sp. Female - From horse, Okunakayama, Iwate Pref., Japan, 16 October 1938, Yajima's collection. Reproduced from Keegan & Toshioka (1957), pi. 12.

27 Dermaccntor sp. Male - Same as above. Reproduced from Keegan & Toshioka ( 1957), pi. 13.

28 Dermaccntor sp. 65 Male from 406 MGL Coll. #101 - From Wild pig, Shizuhara, Kyoto Pref., Japan, 12

December 1955, Akiyama. Reproduced from Keegan & Toshioka ( 1957), pi. 14.

29 Haemaphysalis campanulata Female from Coll. #52-J-004 - From dog, Miura Peninsula, Kanagawa Pref., Japan,

Asanuma. Reproduced from Keegan & Toshioka ( 1957), pi. 17.

30 Haemaphysalis campanulata Male from Coll.#41-J-002 - From Man, Takarazuka, Hyogo Pref., Japan, 15 January 1941, Hiromatsu. Reproduced from Keegan & Toshioka (1957), pi. 18.

31 Haemaphysalis campanulata Nymph from Coll. #67-J-0 188 - From dog, Chiba Pref, Japan, 12 June 1967.

32 Haemaphysalis campanulata Larva from Coll.#67-J-01SS-F| - Reared from material m Coll. #67-J-0 188.

33 Haemaphysalis coneinna [emale - From cow, .'\tsukeshi, Hokkaido, Japan, July 1966, Kitaoka's collection.

34 Haemaphysalis coneinna Male - Nozukesaki. Nemuro-shi, Hokkaido, Japan, May 1964, Kitaoka's collection.

35 Haemaphysalis coneinna Nymph - Reared specimen loaned by Kitaoka.

36 Haemaphysalis coneinna Larva - Same as above.

37 Haemaphysalis sp. (W. Female - From cow, Hachijo (Is.), Tokyct, Japan, 29 May 1949, Asanuma's collec- cornigera group) tion. Reproduced from Keegan & Toshioka (1957), pi. 19.

- pi. 38 Haemaphysalis sp. (//. Male Same as above. Reproduced from Keegan & Toshioka ( 195 7), 20. cornigera group) 39 Haemaphysalis sp. (//. Nymph - Reared specimen loaned by Kitaoka. cornigera group) 40 Haemaphysalis sp. (//. Larva from CoU. #67-J-0?06-F| - From cow, Hachijo Koshima, Tokyo. Japan. 13 cornigera group) July 1967, Yamaguti.

41 Haemaphysalis doenitzi Female from Coll. #56-J-057 - From pheasant, Akune, Kagoshima Pref., Japan, 10

January 1956, Toshioka. Reproduced from Keegan & Toshioka ( 1957), pi. 28.

42 Haemaphysalis flava Female from Coll. #55-J-018 - I'rom dog, Yamakita-cho. Kanagawa Pref., Japan, 4

November 1955. Akiyama. Reproduced from Keegan & Toshioka ( 1957), pi. 21.

4 3 Haemaphysalis /la va Male from Coll. ff55-J-018 - Same as above. Reproduced from Keegan & Toshioka (1957). pi. 22.

44 Haemaphysalis flava Nymph from Coll.#67-J-0294-F| - Reared from material in Coll.#67-J-0294.

45 Haemaphysalis flava Larva from Coll.#67-J-0294-F'| - Same as above.

46 Haemaphysalis formosensis Female from Coll. #67-J-0216 - From Sus scrofa riukiuanus (Wild pig), LIkenson, Amaini Oshima, Kagoshima Pref., Japan, 18 March 1967, Tipton & Mizusawa.

47 Haemaphysalis formosensis Male from Coll. #67-J-021 7 - On vegetation. Ukenson. Amami Oshima, Kagoshima Pref., Japan, 23 March 1967. Tipton & Mizusawa.

48 Haemaphysalis formosensis Nymph from Coll.#67-J-O2l6-1', - Reared from material in Coll.#67-J-()216.

49 Haemaphysalis formosensis Larva from Coll. #67-J-0216-l'| - Reared tfom material in Coll. #67-J-0216.

50 Haemaphysalis fujisana Female from cow, foot of Mt. I uji, near I ujinonin.i. Slii/uoka. Illustrations from Kitoaka (1970).

5 1 Haemaphysalis fujisana Male from cow. foot of Mt. I u|i. near 1 ujinonuya, Shi/uoka. Illusiralions from Kitoaka (1970). 1

Bioi.ocicAi, Skkies, Vol, 15, No. Ticks 223

52 Hacinaphysalis fujisana Nymph Iroin cow, foot of Mt. luji, near lujlnoiniya, Sliizuoka. Illustrations from Kitoaka(1970).

5 3 ilacinaphysalis fujisana Larva from cow, foot of Mt. Fuji, near Fnijinomiya, Shi/uoka. Illustrations from Kitoaka(1970).

- 54 Hacmaphysalis hystricis Female from 406 MGL Coll. #92 From Wild pig, mountain near Inoda, Ishigaki (Is.), Ryukyu Islands, 8 November 1955, Keegan & Toshioka. Reproduced from Keegan & Toshioka (1957). pi. 23.

55 Hacmaphysalis hystricis Male from 406 MGL Coll. #92 - Same as above. Reproduced from Keegan & Toshioka (1957), pi. 24.

- 5 b Hacmaphysalis hystricis Nymph frorti Coll. #67-1-007 I From Sus scrofa riukiuanus (Wild pig), Yamatoson, Amami Oshima, Kagoshima Pref., 21 March 1967, Mizusawa.

57 Hacmaphysalis hystricis Larva from Coll. #67-1-007 1-F, - Reared from material in Coll. #67-1-0071.

5 8 Hacmaphysalis japonica Female from Coll. #67-1-0198 - From cow, Kuttari, Tokachi, Hokkaido, lapan, 13 May 1966, Taniguchi.

5 9 Hacmaphysalis japonica Male - From Capricornis crispus (Serow), Shimajima, Nagano Pref., Japan, 29 November 1953. Sakaguchi. Reproduced from Keegan & Toshioka (1957), pi. 27.

60 Hacmaphysalis japonica Nymph - Same as the lot of CoU. #67-1-0198.

6 Hacmaphysalis kitaokai Female from Coll. #54-1-006 - From Cervus nippon nippon (Sika Deer), Ouchiyama, Mie Pref.. lapan, 17 January 1954, Asanuma. Reproduced from Keegan & Toshioka (1957). pi. 25.

62 Hacmaphysalis kitaokai Male from Coll. #54-1-006 - Same as above. Reproduced from Keegan & Toshioka (1957), pL 26.

63 Hacmaphysalis kitaokai Nymph from Coll.#67-1-0286-F, - Reared from material in Coll. #67-1-0286.

64 Hacmaphysalis kitaokai Larva from Coll.#67-1-0286-F, - Same as above.

- Hakodate, Hokkaido, 65 Hacmaphysalis longicornis Female from Coll. #55-1-049 From horse, pasture near Japan, 17 August 1955, Toshioka & Akiyama. Reproduced from Keegan & Toshioka (1957), pi. 15.

- pasture, Hiroshima Pref., 66 Hacmaphysalis longicornis Male from CoU. #55-1-009 From cow, Azumayama Japan, 19 September 1955, Toshioka & Akiyama. Reproduced from Keegan & Toshioka (1957), pi. 16.

- Abashiri, Hokkaido, Japan, 67 Hacmaphysalis longicornis Nymph from Coll. #67-1-0191 On vegetation. Misaki, 13 July 1967, Toshioka & Sasagawa.

68 Hacmaphysalis longicornis Larva from Coll.#66-J-0022-Fi - Reared from material in Coll.#66-J-0022.

69 Hacmaphysalis mcgaspinosa Female from Coll,#68-J-0301-F, - Reared from material in Coll.#68-J-0301.

70 Hacmaphysalis mcgaspinosa Male from Coll. #68-1-030 l-F, - Reared from material in Coll. #68-1-0301.

- as above. 1 1 Hacmaphysalis mcgaspinosa Nymph from Coll.#68-J-0301-l-, Same

11 Hacmaphysalis mcgaspinosa Larva - Reared specimen loaned by Kitaoka.

73 HacmaphysaUs pcnlalagi Female from Coll. #68-J-0315 - From Pcntalagiis furnessi (Ryukyu Rabbit), Yuwandake, Amami Oshima, Kagoshima Pref.. lapan, 6 April 1968, Mizusawa.

74 HacmaphysaUs pentalagi Male from Coll. #68-1-03 15 - Same as above.

75 Haemaphysalis pentalagi Nymph from Coll. #68-1-031 1-F, - Reared from material in CoU. #68-1-03 1 1.

76 Haemaphysalis pentalagi Larva from Coll. #68-1-031 1-F, - Same as above.

11 HacmaphysaUs wcllingtoni Male - Adopted from Asanuma & Kosaka (1954). pp. 105-106. figs. I & 2.

- Mt. Gomadan, Wakayama Pref., 1 8 Ixodes acu titarsu s Female from RML Coll. #27491 From Man, lapan, 4 luly 1949. Reproduced from Keegan & Toshioka (1957), pi. 29. 1

224

79 Ixodes angiistus

80 Ixodes angiistus

8 Ixodes angustus

8 2 Ixodes gran u la tu s

83 Ixodes gramdatus

84 Ixodes granulatus

85 Ixodes granulatus

86 Ixodes lividus

87 Ixodes lividus

88 Ixodes lividus

89 87521

Biological Series, Vol. 15. No. 1 Ticks 225

August 1955. Toshioka & Akivama. Reproduced from Keegan & Toshioka (1957), pi. 35.

1 05 Ixodes signatus Male from 406 MGL Coll. #40 - Same as above, 14 October 1955. Akivama. Reproduced from Keegan & Toshioka (1957). pi. 36.

1(16 Ixodes signatus Nymph from Coll. #67-1-0092 - Same as above. 12 May 1967, Toshioka & Sasagawa.

117 - 1 Ixodes signatus Larva from Coll. #66-1-0323 Same as above, 15 August 1966, Toshioka & Sasagawa.

1 08 Ixodes simplex simplex Female from RML Coll. #30943 - From Myotis macrodactylus (Little Brown Bat), near Kameaka, Honshu, Japan, 30 September 1952. Reproduced from Keegan & Toshioka (1957), pi. 37.

1 09 Ixodes simplex simplex Nymph - From Myotis macrodactylus (Little Brown Bat), Iwatc Pref.. Japan, 26 July 1967, Yaniaguti (Endo's collection).

110 Ixodes tanuki Female - Adopted from Saito (1964), p. 62, tigs. 1-5.

1 1 I Ixodes tanuki Male - Adopted from Saito (1964). p. 64. figs. 6-9.

1 1 Ixodes turdus Female from Coll. #55-J-024 - From Turdus celaenopus celaenopus (Seven Islands Thrush), Hachijo (Is.), Tokyo, Japan, 12 December 1955, Asanuma. Reproduced from Keegan & Toshioka (1957), pi. 38.

1 1 3 Ixodes turdus Nymph from Coll. #55-1-024 - Same as above.

1 14 Ixodes turdus Larva - Adopted from Asanuma & Sekikawa ( 1952). p. 112, fig. 5.

1 1 Ixodes vespertilionis Female from Coll. #56-1-037 - From Rhinolophus ferrum-equinum nippon (Horse- shoe Bat), Izuru, Tochigi Pref., Japan, 24 February 1956, Toshioka & Akiyama. Reproduced from Keegan & Toshioka (1957), pi. 39.

- 1 I 6 Ixodes vespertilionis Male from 406 MGL Coll. #191 Same as above. Moulted from a nymph of the lot above. Reproduced from Keegan &. Toshioka (1957). pi. 40.

1 1 Ixodes vespertilionis Nymph from Coll. #66-1-0015 - From Rhinolophus cornutus cornutus (Horseshoe Bat), Iwanai, Shiribeshi, Hokkaido, Japan, 7 May 1965, Hattori.

- 1 1 Ixodes vespertilionis Larva from Col). #66-J-0016 Same as above.

1 19 Rhipicephalus sanguineus Female from Coll. #66-R-0005 - From dog, Okinawa. Ryukyu Islands. 19 October 1966. Doucette & Sugiyama.

120 Rhipieephalus sanguineus Male from Coll. #64-1-0279 - From dog. Yokohama-shi. Kanagawa Pref.. Japan, 7 October 1964, Flint.

1 2 Rhipicephalus sanguineus Nymph - Reared specimen sent from Texas, USA.

1 22 Rhipicephalus sanguineus Larva - Same as above. 226 BUIGHAM ^OUNG UNIVERSITY SciENCE BULLETIN

APPENDIX 5

Corrected Scientific Names of Hosts Recorded in the Literature

Some of the scientific names of hosts cited in the Htera- Keisuke Kobayashi (1965). Birds of Japan in natural colors, ture require revision in accord with current knowledge of Hoikusha Publ. Co. Ltd., Osaka, 231 p. Rev. and Knl. Ed. vertebrate taxonomy. Therefore, corrected names were used The Ornithological Society of Japan (1958). A handlist of in Appendix 2. Revisions were based on the following pub- the Japanese birds, 4th Rev. Ed. lications; The English common names were taken trom the follow- Yoshinori Imai/umi (1960). Colored illustrations of the ing publication:

mammalsof Japan. Hoikusha Publ. Co. Ltd., Osaka. 196 p. Ernest P. Walker ( 1964). Mammals of the world. Vols. 1 & 2. Kenji Nakamura & Shun-ichi Ueno (1963). Japanese reptiles and amphibians in color, Hoikusha Publ. Co. Ltd.. Osaka,

214 p.

Page in Name cited Corrected Name ("ommon Name Liter ature ^___ Appendix 2

Apodemus geisha Apodemus argenteus lield mouse Asanumu & Sekikawa ( 1952). (1953) 210,212.214

Apodemtis sylvaticus ainu Apodemus speciosus ainu I ield mouse Asanuma & Sekikawa ( 1952) 207.208.210. 211,214.

Apodemus sylvaticus Apodemus speciosus licid mouse Asanuma & Sekikawa ( 1952) 197.209,216 speciosus Cervus nippon nippon Cervus nippon Sika deer Hoogstraa! (1969) 189. 195, Crocidura russula Crocidura dsinezumi Shrew Sailo (1959b). (1964). Saito et al. 206. 2 in. (1965) 212. 214 Cvclemys Jlavnmar^inala C 'yclemys fla vo mar^ina la Turtle Keegan & Toshioka (1957) Jlavomarginafa

Hori'iiiS canlans cantans Cctlta diphonc canlans Japanese Bush Asanuma & Kusaka ( 1955) Warbler

Horeires canlans ijimae Cciiia diphonc ijimac Japanese Bush Avinunia & Kos.ika ( 1955) Warbler

l.xos amauroiis amauroiis Hvpsipclcs amauroiis Brown-cared Asanunu & Kosaka i 195 5). Asanuma & amauroiis Biilbul Nakagawa (1955) l.epus linudus an^usfidens I.cpus brachyurus angus- Hare Saitol 1959b). Saito et al. (1965),

tidens kilaoka & Saitu ( 1967). Asanuma,

Sakurai et al. (1955)

I.cpus timidus brachyurua I.cpus brachyurus Hare Asanuma ( 1956)

Lepus tinudus lyoni l.epus brachyurus lyoni Hare Saito ( 1959b). Saito et al. ( 1965) Meles anakuma Meles melcs anakuma Badger Saito et al. (1965)

Microhyla Jissipcs Microhyla ornata I rog Sugimoto (1937a) Nemorrhacdus cnspus Capricorms crispus Japanese Serow Neumann (1906). (1911). Nutta!l&

cnspwi \V,irlHirlon ( 1915)

Phasianus versicolor Phasianus colchicus Pheasant Saitu ( 1959b). Saito ct al ( 1965) rabustipcs

al. Phasianus versicolor Phasianus colchicus (jreen Pheasant Asanuma ( 1956). Asanuma, Sakurai et tohkatdt tohkaidi (1955) Phylloscopiis occipi- Phyltoscopus ijimac WilknvWarbler Asanuma and Ktisaka (l'>55) talis ijtmae

Sciurus vulgaris lis Sciurus hs Squirrel Asanuma ( 1956). .Asanuma, Sakurai ct al (1955)

Sus leucomystax or Sus scrofa leucomystax Wild Pig Asanuma, Sakurai et al. ( 1955).' Susl. leucomystax Kishida (1922a)

Syromaticus socmmcrringn Pha stanu s vo cm m crringii Hiindo Copper Asanuma ( 1956). Asanuma. Sakurai et al. scintillans \cinitllans Pheasant (1955)

i'rsus thihctamis /aponicus Sclenarctos thibclanus Asialie Black Keegan & Toshioka ( 1957), Asanuma ( 19561. japomcus Bear Saito (1959b)

PuJJinus Caloncctris Streaked Kenans and Kohls i 1970) leucomelas Icucomclas Shearwater