Individual Species Accounts

Abstract

1 – H. aborensis Warburton, 1913 (Parasitology, 6: 121–130)

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1 – H. aborensis Warburton, 1913 (Parasitology, 6: 121–130)

Type depository: IM (holotype) (Hoogstraal, H., Dhanda, V. & El Kammah, K.M. 1971. Aborphysalis, a new subgenus of Asian Haemaphysalis ticks; and identity, distribution and hosts of H. aborensis Warburton (resurrected) (Ixodoidea: Ixodidae). J. Parasitol., 57: 748–760)

Known stages: male, female, nymph, larva

Zoogeographic Region: Oriental

Ecoregion: northern Indochina subtropical forest

Hosts: Artiodactyla: Bovidae, Suidae, Moschidae; Carnivora: Mustelidae; Rodentia: Hystricidae (AN)

Artiodactyla: Cervidae; Carnivora: Felidae (A)

Galliformes: Phasianidae (NL)

Scandentia: Tupaiidae; Passeriformes: Paridae (N)

Human infestation: no

Remarks: Mitchell (1979) listed Muridae among the alleged hosts of Nepalese ticks collected between 1966 and 1970, including H. aborensis (stage not stated). However, in Hoogstraal’s (1971) resurrection of H. aborensis, murids are not mentioned as hosts, and for that reason they do not appear in our host list.

References

Chen, Z., Yang, X., Bu, F., Yang, X., Yang, X. & Liu, J. 2010. Ticks (Acari: Ixodoidea: Argasidae, Ixodidae) of China. Exp. Appl. Acarol., 51: 393–404.

Hoogstraal, H., Dhanda, V. & El Kammah, K.M. 1971. Aborphysalis, a new subgenus of Asian Haemaphysalis ticks; and identity, distribution and hosts of H. aborensis Warburton (resurrected) (Ixodoidea: Ixodidae). J. Parasitol., 57: 748–760.

Mitchell, R.M. 1979. A list of ectoparasites from Nepalese mammals, collected during the Nepal ectoparasite program. J. Med. Entomol., 16: 227–233.

Phan Trong, C. 1977. Ve bet va con trung ky sinh o Viet Nam. Tap 1. Ve (Ixodoidea), mo ta va phan loai. Ha Noi: Khoa hoc va ky thuat, 489 pp. In Vietnamese.

Xu, R. & Li, K. 1997. A collection of ticks from Guizhou, China. Syst. Appl. Acarol., 2: 245–246.

2 – H. aciculifer Warburton, 1913 (Parasitology, 6: 121–130)

Type depository: BMNH (lectotype, paralectotypes) (Keirans, J.E. & Hillyard, P.D. 2001. A catalogue of the type specimens of Ixodida (Acari: Argasidae, Ixodidae) deposited in The Natural History Museum, London. Occ. Pap. Syst. Entomol. (13), 74 pp.)

Known stages: male, female, nymph, larva

Zoogeographic Region: Afrotropical

Ecoregions: tropical and subtropical grasslands, savannas and shrublands; tropical and subtropical moist broadleaf forests

Hosts: usual hosts for adult ticks are Artiodactyla: Bovidae.

Artiodactyla: Bovidae; Carnivora: Viverridae; Lagomorpha: Leporidae (ANL)

Carnivora: Canidae, Herpestidae (AL)

Artiodactyla: Suidae; Carnivora: Felidae (A)

Rodentia: Muridae; Galliformes: Phasianidae (NL)

Erinaceomorpha: Erinaceidae; Passeriformes: Corvidae (N)

Carnivora: Hyaenidae; Rodentia: Cricetidae (L)

Human infestation: no

Remarks: data concerning H. aciculifer prior to Hoogstraal and El Kammah (1972) are ignored here because of confusion with related tick species that made earlier diagnoses uncertain. See also H. rugosa.

References

Hoogstraal, H. & El Kammah, K.M. 1972. Notes on African Haemaphysalis ticks. X. H. (Kaiseriana) aciculifer Warburton and H. (K.) rugosa Santos Dias, the African representatives of the spinigera subgroup (Ixodoidea: Ixodidae). J. Parasitol., 58: 960–978.

Horak, I.G. & Boomker, J. 1998. Parasites of domestic and wild animals in South Africa. XXXV. Ixodid ticks and bot fly larvae in the Bontebok National Park. Onderstepoort J. Vet. Res., 65: 205–211.

Horak, I.G., Fourie, L.J. & Boomker, J. 1997. A ten-year study of ixodid tick infestations of bontebok and grey rhebok in the Western Cape Province, South Africa. S. Afr. J. Wildl. Res., 27: 5–10.

Norval, R.A.I. 1985. The ticks of Zimbabwe. XIV. The lesser known Haemaphysalis species. Zimbabwe Vet. J., 16: 54–59.

Walker, J.B. 1974. The ixodid ticks of Kenya. A review of present knowledge of their hosts and distribution. Commonwealth Institute of Entomology, London, 220 pp.

3 – H. aculeata Lavarra, 1904 (Bol. Soc. Zool. Ital. Ser. 2, 5: 255–258)

Type depository: BMNH (lectotype) (Keirans and Hillyard 2001, op. cit. under H. aciculifer)

Known stages: male, female, nymph, larva

Zoogeographic Region: Oriental

Ecoregions: tropical and subtropical dry broadleaf forests; desert and xeric shrublands

Hosts: usual hosts for larvae, nymphs and adults are Artiodactyla: Tragulidae.

Artiodactyla: Tragulidae (ANL)

Artiodactyla: Bovidae (AN)

Artiodactyla: Cervidae; Carnivora: Felidae, Herpestidae; Primates (unknown family); Aves (unknown order) (A)

Primates: Cercopithecidae; Galliformes: Phasianidae (NL)

Rodentia: Hystricidae; Cuculiformes: Cuculidae; Passeriformes: Pycnonotidae, Sturnidae, Timaliidae (N)

Rodentia: Muridae; Passeriformes: Muscicapidae (L)

Human infestation: yes (Keirans 1985)

Remarks: all parasitic stages of H. aculeata were redescribed by Geevarghese et al. (2009). Without further explanation, Geevarghese et al. (1997) reduced the host range of this tick to “man, monkey and birds.” We consider records from other types of hosts in the references below to be provisionally valid.

References

Bhat, H.R. & Sreenivasan, M.A. 1981. Further records of the ticks of some reptilian and mammalian hosts in the Kyasanur Forest disease area, Karnataka, India. Ind. J. Parasitol., 5: 207–210.

Geevarghese, G., Mandke, O.A. & Mishra, A.C. 2009. Haemaphysalis (Kaiseriana) aculeata Lavarra, 1904 (Ixodoidea: Ixodidae) re-description of adult and immature stages. Acarologia, 49: 5–11.

Geevarghese, G., Fernandes, S. & Kulkarni, S.M. 1997. A checklist of Indian ticks (Acari: Ixodoidea). Ind. J. Anim. Sci., 67: 566–574.

Keirans, J.E. 1985. George Henry Falkiner Nuttall and the Nuttall tick catalogue. U. S. Dept. Agric., Agric. Res. Ser. Misc. Pub. (1438), 1785 pp.

Rajagopalan, P.K. 1972. Ixodid ticks (Acarina: Ixodidae) parasitizing wild birds in the Kyasanur Forest disease area of Shimoga District, Mysore State, India. J. Bombay Nat. Hist. Soc., 69: 55–78.

Rajagopalan, P.K. & Sreenivasan, M.A. 1981. Ixodid ticks on cattle and buffaloes in the Kyasanur Forest disease area of Karnataka State. Ind. J. Med. Res., 73: 880–889.

Rajagopalan, P.K., Patil, A.P. & Boshell, J. 1968. Ixodid ticks on their mammalian hosts in the Kyasanur Forest disease area of Mysore State, India, 1961–1964. Ind. J. Med. Res., 56:510–526.

Seneviratna, P. 1965. The Ixodoidea (ticks) of Ceylon. Parts II and III. Ceylon Vet. J., 13: 28–54.

Trapido, H., Goverdhan, M.K., Rajagopalan, P.K. & Rebello, M.J. 1964. Ticks ectoparasitic on monkeys in the Kyasanur Forest disease area of Shimoga District, Mysore State, India. Am. J. Trop. Med. Hyg., 13: 763–772.

4 – H. adleri Feldman-Musham, 1951 (Bull. Res. Counc. Israel Ser. C, 1: 96–107)

Type depository: not stated (Feldman-Muhsam, B. 1951. A note on east Mediterranean species of the genus Haemaphysalis. Bull. Res. Counc. Israel Ser. C, 1: 96–107)

Known stages: male, female

Zoogeographic Region: Palearctic

Ecoregions: desert and xeric shrublands

Hosts: Carnivora: Mustelidae (AN)

Artiodactyla: Suidae; Carnivora: Canidae, Felidae (A)

Human infestation: no

Remarks: Kolonin (2009) does not include hosts for nymphs of H. adleri, probably because this stage has not been described. We provisionally accept the record of Theodor and Costa (1967) of nymphs of this tick on Mustelidae.

References

Keysary, A., Eremeeva, M.E., Leitner, M., Din, A.B., Wikswo, M.E., Mumcuoglu, K.Y., Inbar, M., Wallach, A.D., Shanas, U., King, R. & Waner, T. 2011. Spotted fever group rickettsiae collected from wild animals in Israel. Am. J. Trop. Med. Hyg., 85: 919–923.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Morel, P.-C. 2003. Les tiques d’Afrique et du Bassin méditerranéen (1965–1995). CIRAD- EMVT, 1342 pp.

Theodor, O. & Costa, M. 1967. A survey of the parasites of wild mammals and birds in Israel. Part one. Ectoparasites. The Israel Academy of Science and Humanities, Jerusalem, 119 pp.

5 – H. anomala Warburton, 1913 (Parasitology, 6: 121–130)

Type depository: IM (holotype) (Hoogstraal, H., Kohls, G.M. & Trapido, H. 1967. Studies on Southeast Asian Haemaphysalis ticks (Ixodoidea, Ixodidae). H. (Kaiseriana) anomala Warburton: redescription, hosts, and distribution. J. Parasitol., 53: 196–201) as Haemaphysalis cornigera anomala

Known stages: male, female, nymph, larva

Zoogeographic Region: Oriental

Ecoregions: tropical and subtropical moist broadleaf forests

Hosts: usual hosts for adult ticks are Artiodactyla: Bovidae.

Artiodactyla: Bovidae, Cervidae; Carnivora: Canidae, Felidae (A)

Rodentia: Muridae; Cuculiformes: Cuculidae (NL)

Galliformes: Phasianidae; Passeriformes: Turdidae (stages unknown)

Human infestation: yes (Tanskul et al. 1983)

References

Hoogstraal, H., Dhanda, V. & Bhat, H.R. 1972. Haemaphysalis (Kaiseriana) anomala Warburton (Ixodoidea: Ixodidae) from India: description of immature stages and biological observations. J. Parasitol., 58: 605–610.

Mitchell, R.M. & Dick, J.A. 1978. Ectoparasites from Nepal birds. J. Bombay Nat. Hist. Soc., 74: 264–274.

Tanskul, P., Stark, H.E. & Inlao, I. 1983. A checklist of ticks of Thailand (Acari: Metastigmata: Ixodoidea). J. Med. Entomol., 20: 330–341.

6 – H. anomaloceraea Teng, 1984 (In Teng and Cui, 1984, Acta Zootax. Sin., 9: 37–40. In Chinese)

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. (1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.) regard this species as a synonym of H. shimoga without justifying their decision. Kolonin, G.V. (2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/) considers H. anomaloceraea and also H. shimoga to be synonyms of H. taiwana but does not elaborate. We follow Guglielmone, A.A., Robbins, R.G., Apanaskevich, D.A., Petney, T.N., Estrada-Peña, A., Horak, I.G., Shao, R. & Barker, S.C. (2010. The Argasidae, Ixodidae and Nuttalliellidae (Acari: Ixodida) of the world: a list of valid species names. Zootaxa, 2528: 1–28), who regard H. anomaloceraea as a valid species, pending a robust justification of the positions taken by the authors cited above.

Type depository: IZAS (holotype) (Teng, K.-F. & Cui, Y.-Q. 1984. Descriptions of a new species of Haemaphysalis and male of H. primitiva Teng, 1982 from Yunnan. Acta Zootax. Sin., 9: 37–40. In Chinese)

Known stages: male

Zoogeographic Region: Oriental

Ecoregions: Nujiang Langcang Gorge alpine conifer and mixed forests

Hosts: unknown

Human infestation: no

Remarks: see above.

References

Chen, Z., Yang, X., Bu, F., Yang, X., Yang, X. & Liu, J. 2010. Ticks (Acari: Ixodoidea: Argasidae, Ixodidae) of China. Exp. Appl. Acarol., 51: 393–404.

Teng, K.-F. & Cui, Y.-Q. 1984. Descriptions of a new species of Haemaphysalis and male of H. primitiva Teng, 1982. Acta Zootax. Sin., 9: 37–40.

7 – H. anoplos Hoogstraal, Uilenberg & Klein, 1967 (J. Parasitol., 53: 1103–1105)

Type depositories: USNTC (holotype), HH (paratype) (Hoogstraal, H., Uilenberg, G. & Klein, J.-L. 1967. Haemaphysalis (Rhipistoma) anoplos sp. n., a spurless tick of the elongata group (Ixodoidea, Ixodidae) parasitizing Nesomys rufus Peters (Rodentia) in Madagascar. J. Parasitol., 53: 1103–1105)

Known stages: female

Zoogeographic Region: Afrotropical

Ecoregion: Madagascar subhumid forests

Hosts: Rodentia: Nesomyidae (A)

Human infestation: no

Reference

Uilenberg, G., Hoogstraal, H. & Klein, J.-M. 1979. Les tiques (Ixodoidea) de Madagascar et leur rôle vecteur. Arch. Inst. Pasteur Madagascar Num. Spéc., 153 pp.

8 – H. aponommoides Warburton, 1913 (Parasitology, 6: 121–130)

Type depository: BMNH (lectotype, paralectotypes) (Keirans and Hillyard 2001, op. cit. under H. aciculifer) originally named Haemaphysalis inermis aponommoides

Known stages: male, female, nymph, larva

Zoogeographic Regions: Oriental, Palearctic

Ecoregions: tropical and subtropical grasslands, savannas and shrublands; temperate conifer forests

Hosts: usual hosts for adult ticks are Artiodactyla: Bovidae.

Artiodactyla: Bovidae, Cervidae; Perissodactyla: Equidae; Carnivora: Canidae, Felidae, Ursidae; Rodentia: Sciuridae (A)

Rodentia: Muridae; Soricomorpha: Soricidae; Galliformes: Phasianidae (NL)

Human infestation: yes (Hoogstraal and Mitchell 1971)

Remarks: Mitchell (1979) listed all hosts of Nepalese ticks collected between 1966 and 1970, including H. aponommoides (no developmental stage stated), which was allegedly found on Cricetidae. However, cricetids are not included by Hoogstraal and Mitchell (1971) in their study of this tick, and consequently this host record does not appear in our list of hosts of H. aponommoides.

References

Chen, Z., Yang, X., Bu, F., Yang, X. & Liu, J. 2010. The ticks (Acari: Ixodoidea: Argasidae, Ixodidae) of China. Exp. Appl. Acarol., 51: 393–404.

Hoogstraal, H. & Mitchell, R.M. 1971. Haemaphysalis (Alloceraea) aponommoides Warburton (Ixodoidea: Ixodidae), description of immature stages, hosts, distribution, and ecology in India, Nepal, Sikkim, and China. J. Parasitol., 57: 635–645.

Mitchell, R.M. 1979. A list of ectoparasites from Nepalese mammals, collected during the Nepal ectoparasite program. J. Med. Entomol., 16: 227–233.

Phan Trong, C. 1977. Ve bet va con trung ky sinh o Viet Nam. Tap 1. Ve (Ixodoidea), mo ta va phan loai. Ha Noi: Khoa hoc va ky thuat, 489 pp. In Vietnamese.

9 – H. asiatica (Supino, 1897) (Atti Soc. Veneto-Trentina Sci. Nat. Residente Padova, Ser. 2, 3: 230–238)

Type depository: GM (holotype) (Hoogstraal, H. & Trapido H. 1966. Studies on Southeast Asian Haemaphysalis ticks (Ixodoidea, Ixodidae). Species described by Supino in 1897 from Burma, with special reference to H. (Rhipistoma) asiaticus (= H. dentipalpis Warburton and Nuttall). J. Parasitol., 52: 1172–1187) as Opisthodon asiaticus. See also below.

Known stages: male, female, nymph, larva

Zoogeographic Region: Oriental

Ecoregions: tropical and subtropical forests

Hosts: usual hosts for adult ticks are Carnivora: Viverridae.

Scandentia: Tupaiidae (ANL)

Carnivora: Canidae, Felidae (AN)

Artiodactyla: Suidae; Carnivora: Viverridae (A)

Rodentia: Muridae (NL)

Rodentia: Sciuridae (N)

Human infestation: no

Remarks: this tick has been treated as a synonym of H. leachi by several authors, but H. asiatica is a legitimate species, as stated in Hoogstraal and Trapido (1966), who refer to this tick as Haemaphysalis asiaticus. Phan Trong (1977) found adults of H. asiatica on Aves (Cuculiformes and Passeriformes), but we feel that these records need confirmation and they are not included in our host list. Kolonin (2009) does not recognize Scandentia as hosts of adults, but we consider this relationship, cited by Tanskul et al. (1983), to be valid.

References

Chen, Z., Yang, X., Bu, F., Yang, X., Yang, X. & Liu, J. 2010. Ticks (Acari: Ixodoidea: Argasidae, Ixodidae) of China. Exp. Appl. Acarol., 51: 393–404.

Grassman, L.I., Sarataphan, N., Tewes, M.E., Silvy, N.J. & Nakanakrat, T. 2004. Ticks (Acari: Ixodidae) parasitizing wild carnivores in Phu Khieo wildlife sanctuary. J. Parasitol., 90: 657–659.

Hoogstraal, H. & Trapido, H. 1966. Studies on Southeast Asian Haemaphysalis ticks (Ixodoidea: Ixodidae). Species described by Supino in 1897 from Burma, with special reference to H. (R.) asiaticus (= H. dentipalpis Warburton and Nuttall). J. Parasitol., 52: 1172–1187.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Parola, P., Cornet, J.-P., Sanogo, Y.O., Miller, R.S., Thien, H.V., González, J.-P., Raoult, D., Telford, S.R. & Wongsrichanalai, C. 2003. Detection of Ehrlichia spp., Anaplasma spp. Rickettsia spp., and other eubacteria in ticks from the Thai-Myanmar border and Vietnam. J. Clin. Microbiol., 41: 1600–1608.

Phan Trong, C. 1977. Ve bet va con trung ky sinh o Viet Nam. Tap 1. Ve (Ixodoidea), mo ta va phan loai. Ha Noi: Khoa hoc va ky thuat, 489 pp. In Vietnamese.

Saito, Y., Hoogstraal, H. & Vasuvat, C. 1971. Studies on Southeast Asian Haemaphysalis ticks (Ixodoidea: Ixodidae). Identity and hosts of immature stages of H. (Rhipistoma) asiatica (Supino) from Thailand. J. Parasitol., 57: 1099–1103.

Tanskul, P., Stark, H.E. & Inlao, I. 1983. A checklist of ticks of Thailand (Acari: Metastigmata: Ixodoidea). J. Med. Entomol., 20: 330–341.

Xu, R. & Li, K. 1997. A collection of ticks from Guizhou, China. Syst. Appl. Acarol., 2: 245–246.

10 – H. atheruri Hoogstraal, Trapido & Kohls, 1965 (J. Parasitol., 51: 114–125)

Type depositories: USNTC (holotype, paratypes), BMNH, IMRKL (paratypes) (Keirans and Hillyard 2001, op. cit. under H. aciculifer) as Haemaphysalis atherurus

Known stages: male, female, nymph, larva

Zoogeographic Region: Oriental

Ecoregions: tropical and subtropical forests

Hosts: usual hosts for larvae, nymphs and adults are Rodentia: Hystricidae.

Rodentia: Hystricidae (ANL)

Scandentia: Tupaiidae; Carnivora: Canidae (A)

Human infestation: no

Remarks: see H. traguli, which has been confused with H. atheruri.

References

Hoogstraal, H., Trapido, H. & Kohls, G.M. 1965. Studies on Southeast Asian Haemaphysalis ticks (Ixodoidea, Ixodidae). H. atherurus sp. n. and redescription of type material of H. birmaniae Supino, 1897. J. Parasitol., 51: 114–125.

Tanskul, P., Stark, H.E. & Inlao, I. 1983. A checklist of ticks of Thailand (Acari: Metastigmata: Ixodoidea). J. Med. Entomol., 20: 330–341.

11 – H. bancrofti Nuttall & Warburton, 1915 (Ticks. A monograph of the Ixodoidea. Part III. The genus Haemaphysalis. Cambridge University Press, London, pp. 349–550)

See below and also H. novaeguineae.

Type depository: BMNH (lectotype, paralectotypes) (Keirans and Hillyard 2001, op. cit. under H. aciculifer)

Known stages: male, female, nymph, larva

Zoogeographic Regions: Australasian, Oriental

Ecoregions: tropical and subtropical moist broadleaf forests; tropical and subtropical grasslands, savannas and shrublands

Hosts: usual hosts for larvae, nymphs and adults are Diprotodontia: Macropodidae. Aves are considered exceptional hosts for this tick.

Diprotodontia: Macropodidae, Phascolarctidae (ANL)

Artiodactyla: Bovidae; Carnivora: Canidae; Peramelemorphia: Peramelidae (AN)

Dasyuromorphia: Dasyuridae (AL)

Artiodactyla: Suidae; Perissodactyla: Equidae (A)

Diprotodontia: Phalangeridae, Potoroidae; Cuculiformes: Cuculidae (N)

Chiroptera: Vespertilionidae (L)

Artiodactyla: Cervidae; Carnivora: Felidae; Diprotodontia: Macropodidae, Petauridae, Vombatidae; Peramelemorphia: Thylacomyidae; Rodentia: Muridae; Passeriformes: Corvidae, Cracticidae (stages unknown)

Human infestation: yes (Laan et al. 2011)

Remarks: Camicas et al. (1998) state that H. bancrofti is found exclusively in the Australasian Region, but there is a record for the Oriental Region in Hoogstraal and Kim (1985). Kolonin (2009) does not include Aves as hosts of H. bancrofti, but we accept the records in Roberts (1963), Laan et al. (2011) and Owen (2011) from birds. It should be noted, however, that Roberts (1963) states that a female specimen had been found on Cuculidae when, in fact, it was a nymph of H. bancrofti that subsequently molted to the female stage in the laboratory, as clarified by Wilkinson and Utech (1962).

References

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. 1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.

Heath, A.C.G. 1986. Aspects of the biology, seasonality and host associations of Haemaphysalis bancrofti, H. humerosa, H. bremneri and Ixodes tasmani (Acari: Ixodidae). In C. Cremin, C. Dobson & D.E. Moorhouse (editors), Parasite lives. University of Queensland Press, St Lucia, Australia, pp. 179–188.

Hoogstraal, H. & Kim, K.C. 1985. Tick and mammal coevolution, with emphasis on Haemaphysalis. In K.C. Kim (editor), Coevolution of parasitic arthropods and mammals. John Wiley & Sons, New York, pp. 505–568.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Laan, B., Handasyde, K. & Beveridge, I. 2011. Occurrence of the tick Haemaphysalis bancrofti Nuttall & Warburton, 1915 in Victoria with additional data on its distribution and with scanning electron micrographs of life cycle stages. Proc. R. Soc. Victoria, 123: 187–197.

Oakwood, M. & Spratt, D.M. 2000. Parasites of the northern quoll, Dasyurus hallucatus (Marsupialia: Dasyuridae) in tropical savanna, Northern Territory. Aust. J. Zool., 48: 79–90.

Owen, I.L. 2011. Parasites of animals in Papua New Guinea recorded at the National Veterinary Laboratory: a catalogue, historical review and zoogeographical affiliations. Zootaxa, 3143: 1–163.

Roberts, F.H.S. 1963. A systematic study of the Australian species of the genus Haemaphysalis Koch (Acarina: Ixodidae). Aust. J. Zool., 11: 35–80.

Roberts, F.H.S. 1970. Australian ticks. CSIRO, Melbourne, 267 pp.

Wilkinson, P.R. & Utech, K.B.W. 1962. Bird hosts of Haemaphysalis ticks in Australia. Aust. J. Sci., 25: 169–170.

12 – H. bandicota Hoogstraal & Kohls, 1965 (J. Parasitol., 51: 460–466)

Type depositories: USNTC (holotype, paratypes), HH, SEATO (paratypes) (Hoogstraal, H. & Kohls, G.M. 1965. Southeast Asian Haemaphysalis ticks (Ixodoidea, Ixodidae). H. bandicota sp. n. from bandicoot rats in Taiwan, Thailand and Burma. J. Parasitol., 51: 460–466)

Known stages: male, female, nymph, larva

Zoogeographic Region: Oriental

Ecoregions: tropical and subtropical moist broadleaf forests

Hosts: usual hosts for larvae, nymphs and adult ticks are Rodentia: Muridae. Artiodactyla are considered exceptional hosts for this tick.

Rodentia: Muridae (ANL)

Scandentia: Tupaiidae; Carnivora: Herpestidae (A, N and/or L)

Artiodactyla: Bovidae (A)

Human infestation: no

Remarks: Tanskul et al. (1983) use the term “immatures” without specifying which of the sub-adult stages of H. bandicota were found on hosts.

References

Chen, Z., Yang, X., Bu, F., Yang, X., Yang, X. & Liu, J. 2010. Ticks (Acari: Ixodoidea: Argasidae, Ixodidae) of China. Exp. Appl. Acarol., 51: 393–404.

Hoogstraal, H. & Kohls, G.M. 1965. Southeast Asian Haemaphysalis ticks (Ixodoidea, Ixodidae). H. bandicota sp. n. from bandicoot rats in Taiwan, Thailand, and Burma. J. Parasitol., 51: 460–466.

Tanskul, P., Stark, H.E. & Inlao, I. 1983. A checklist of ticks of Thailand (Acari: Metastigmata: Ixodoidea). J. Med. Entomol., 20: 330–341.

13 – H. bartelsi Schulze, 1938 (Z. Morphol. Ökol. Tiere, 34: 135–149)

This species was originally classified as H. koningsbergeri (Wilson, N., Hoogstraal, H. & Kohls, G.M. 1968. Studies on Southeast Asian Haemaphysalis ticks (Ixodoidea, Ixodidae). Redescription of H. (Rhipistoma) bartelsi Schulze (resurrected), the Indonesia flying squirrel haemaphysalid. J. Parasitol., 54: 1223–1227).

Type depository: USNTC (lectotype, paralectotype) (Keirans, J.E. & Clifford, C.M. 1984. A checklist of types of Ixodoidea (Acari) in the collection of the Rocky Mountain Laboratories. J. Med. Entomol., 21: 310–320)

Known stages: male, female, nymph, larva

Zoogeographic Region: Oriental

Ecoregion: Java rain forests

Hosts: Rodentia: Sciuridae (ANL)

Human infestation: no

Reference

Hoogstraal, H., Gaber, S., Van Peenen, P.F.D., Duncan, F. J. & Kadarsan, S. 1972. Haemaphysalis (Rhipistoma) bartelsi Schulze (Ixodoidea: Ixodidae): immature stages from a treehole nest of the Indonesian red giant flying squirrel. J. Parasitol., 58: 989–992.

14 – H. bequaerti Hoogstraal, 1956 (J. Parasitol., 42: 156–172)

Type depositories: USNTC (holotype, paratype), BMNH, HH, OVI (paratypes) (Keirans & Hillyard, 2001, op. cit. under H. aciculifer)

Known stages: male, female, nymph, larva

Zoogeographic Region: Afrotropical

Ecoregions: tropical and subtropical grasslands, savannas and shrublands

Hosts: Hyracoidea: Procaviidae (ANL)

Human infestation: no

Reference

Hoogstraal, H. 1956. Notes on African Haemaphysalis ticks. III. The hyrax parasites, H. bequaerti sp. nov., H. orientalis N. and W., 1915 (new combination), and H. cooleyi Bedford, 1929 (Ixodoidea, Ixodidae). J. Parasitol., 42: 156–172.

15 – H. birmaniae Supino, 1897 (Atti Soc. Veneto-Trentina Sci. Nat. Residente Padova, Ser. 2, 3: 230–238)

Type depositories: GM, BMNH (syntypes) (Keirans and Hillyard 2001, op. cit. under H. aciculifer). See H. darjeeling.

Known stages: male, female, nymph, larva

Zoogeographic Region: Oriental

Ecoregions: temperate broadleaf deciduous forests

Hosts: usual hosts for larvae, nymphs and adults are Artiodactyla: Bovidae and Cervidae.

Artiodactyla: Bovidae, Cervidae (ANL)

Artiodactyla: Suidae; Carnivora: Canidae, Mustelidae; Rodentia: Hystricidae (A)

Human infestation: yes (Hoogstraal 1970)

Remarks: collection data published prior to the work of Hoogstraal (1970) have been ignored because there is considerable uncertainty concerning the identification of H. birmaniae.

References

Chen, Z., Yang, X., Bu, F., Yang, X., Yang, X. & Liu, J. 2010. Ticks (Acari: Ixodoidea: Argasidae, Ixodidae) of China. Exp. Appl. Acarol., 51: 393–404.

Hoogstraal, H. 1970. Haemaphysalis (H.) birmaniae Supino and H. (H.) goral sp. n. (Ixodoidea: Ixodidae), Asian parasites of artiodactyl mammals. J. Parasitol., 56: 1227–1238.

Keirans, J.E. 1985. George Henry Falkiner Nuttall and the Nuttall tick catalogue. U. S. Dept. Agric., Agric. Res. Ser. Misc. Pub. (1438), 1785 pp.

Phan Trong, C. 1977. Ve bet va con trung ky sinh o Viet Nam. Tap 1. Ve (Ixodoidea), mo ta va phan loai. Ha Noi: Khoa hoc va ky thuat, 489 pp. In Vietnamese.

16 – H. bispinosa Neumann, 1897 (Mém. Soc. Zool. Fr., 10: 324–420)

See remarks below.

Type depository: BMNH (neotype) (Keirans and Hillyard 2001, op. cit. under H. aciculifer)

Known stages: male, female, nymph, larva

Zoogeographic Region: Oriental

Ecoregions: tropical and subtropical moist broadleaf forests

Hosts: usual hosts for larvae, nymphs and adults are Artiodactyla: Bovidae and Carnivora: Canidae.

Mammalia (several orders); Galliformes: Phasianidae; Passeriformes: Sturnidae (ANL)

Psittaciformes: Psittacidae (A)

Cuculiformes: Cuculidae; Coraciiformes: Coraciidae; Passeriformes (several families) (NL)

Falconiformes: Accipitridae (L)

Charadriiformes: Charadriidae; Strigiformes: Strigidae (stages unknown)

Human infestation: no

Remarks: many specimens formerly identified as H. bispinosa are in fact H. longicornis, as discussed in Hoogstraal et al. (1968), or several other species of Haemaphysalis (Keirans 1985). Camicas et al. (1998) state that this species is found in the Australasian and Oriental Zoogeographic Regions, and Keirans (1985) presents Afrotropical records for H. bispinosa. We, however, have not found bona fide records for the Australasian Region, and we believe that African records are the result of mislabeling or the introduction of this tick to new environments in which it failed to become established. Guo et al. (2002) state that H. bispinosa has been found in Gansu Province (Palearctic) but this record was not validated in Chen et al. (2010). The latter authors describe the distribution of H. bispinosa as including Hubei Province, whose boundaries overlap the Palearctic and Oriental Regions. Nevertheless, we provisionally regard this species as Oriental. There is a record of H. bispinosa crawling on a human in Audy et al. (1960), but as the tick was not attached, we do not consider humans to be hosts of this species. Other records of H. bispinosa on humans are doubtful or refer to H. longicornis in the Australasian Region. Kolonin (2009) excludes Aves as hosts for adults of H. bispinosa, but the records of Rajagopalan (1972) of males and females on Aves are regarded as valid by us. Dilrukshi (2006) considers it probable that H. bispinosa constitutes a complex of species in Sri Lanka. See also H. longicornis.

References

Arunachalam, K. & Harikrishnan, T.J. 2009. Occurrence of Haemaphysalis bispinosa in a child. Ind. Vet. J., 86: 93.

Audy, J.R., Nadchatram, M. & Lim, B.-L. 1960. Malaysian parasites. XLIX. Host distribution of Malayan ticks (Ixodoidea). Stud. Inst. Med. Res. Malaya, 29: 225–246.

Bhat, H.R. & Sreenivasan, M.A. 1981. Further records of the ticks of some reptilian and mammalian hosts in the Kyasanur Forest disease area, Karnataka, India. Ind. J. Parasitol., 5: 207–210.

Boshell, J. & Rajagopalan, P.K. 1968. Observations on the experimental exposure of monkeys, rodents and shrews to infestation of ticks in forest in Kyasanur Forest disease area. Ind. J. Med. Res., 56 (4) (Suppl.): 573–588.

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. 1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.

Chen, Z., Yang, X., Bu, F., Yang, X., Yang, X. & Liu, J. 2010. Ticks (Acari: Ixodoidea: Argasidae, Ixodidae) of China. Exp. Appl. Acarol., 51: 393–404.

Dilrukshi, P.R.M.P. 2006. Taxonomic status of ticks in Sri Lanka. In C.N.B. Bambaradeniya (editor), The fauna of Sri Lanka: status of taxonomy, research and conservation. The World Conservation Union, Colombo, Sri Lanka, pp. 65–69.

Guo, S., Yuan, Z., Wu, G., Wang, W, Ma, D. & Du, H. 2002. Epidemiology of ovine theileriosis in Ganan Region, Gansu Province, China. Parasitol. Res. (Suppl.), 88: S36-S37.

Hoogstraal, H., Roberts, F.H.S., Kohls, G.M. & Tipton, V.J. 1968. Review of Haemaphysalis (Kaiseriana) longicornis Neumann (resurrected) of Australia, New Zealand, New Caledonia, Fiji, Japan, Korea, and northeastern China and USSR, and its parthenogenetic and bisexual populations (Ixodoidea, Ixodidae). J. Parasitol., 54: 1197–1213.

Hoogstraal, H., Lim, B.-L. & Anastos, G. 1969. Haemaphysalis (Kaiseriana) bispinosa Neumann (Ixodoidea: Ixodidae): evidence for consideration as an introduced species in the Malay Peninsula and Borneo. J. Parasitol., 55: 1075–1077.

Keirans, J.E. 1985. George Henry Falkiner Nuttall and the Nuttall tick catalogue. U. S. Dept. Agric., Agric. Res. Ser. Misc. Pub. (1438), 1785 pp.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Mitchell, R.M. & Dick, J.A. 1978. Ectoparasites from Nepal birds. J. Bombay Nat. Hist. Soc., 74: 264–274.

Rahman, M.H. & Mondal, M.M.H. 1985. Tick fauna of Bangladesh. Ind. J. Parasitol., 9: 145–149.

Rajagopalan, P.K. 1972. Ixodid ticks (Acarina: Ixodidae) parasitizing wild birds in the Kyasanur Forest disease area of Shimoga District, Mysore State, India. J. Bombay Nat. Hist. Soc., 69: 55–78.

Rajagopalan, P.K., Patil, A.P. & Boshell, J. 1968. Ixodid ticks on their mammalian hosts in the Kyasanur Forest disease area of Mysore State, India, 1961–1964. Ind. J. Med. Res., 56: 510–526.

Rebello, M.J. & Reuben, R. 1967. A report on ticks collected from birds and small mammals in North Arcot and Chittoor Districts, South India. J. Bombay Nat. Hist. Soc., 68: 283–289.

Seneviratna, P. 1965. The Ixodoidea (ticks) of Ceylon. Parts II and III. Ceylon Vet. J., 13: 28–54.

Sreenivasan, M.A., Rajagopalan, P.K. & Bhat, H.R. 1972. Isolation of Kyasanur Forest disease virus from ixodid ticks collected between 1965 and 1972. In G.P. Channabasavanna & C.A. Viraktamath (editors), Progress in acarology, volume I. Oxford & IBH Publishing Company, New Delhi, pp. 37–44.

17 – H. borneata Hoogstraal, 1971 (J. Parasitol., 57: 1096–1098)

Type depository: BMNH (holotype) (Keirans and Hillyard 2001, op. cit. under H. aciculifer)

Known stages: male

Zoogeographic Region: Oriental

Ecoregions: tropical and subtropical moist and dry broadleaf forests

Hosts: Artiodactyla: Cervidae (A)

Human infestation: no

Remarks: Mihalca et al. (2011) regard H. borneata as an endangered species.

References

Hoogstraal, H. 1971. Haemaphysalis (Kaiseriana) borneata sp. n. (Ixodoidea: Ixodidae), a tick of the H. (K.) aculeata group parasitizing the sambar deer in Borneo. J. Parasitol., 57: 1096–1098.

Mihalca, A.D., Gherman, C.M. & Cozma, V. 2011. Coendangered hard-ticks: threatened or threatening? Parasit. Vectors, 4 (71), 7 pp.

18 – H. bremneri Roberts, 1963 (Aust. J. Zool., 11: 35–80)

Type depositories: QM (holotype, paratypes), ANIC (paratypes), USNTC (paratypes?). (Roberts, F.H.S. 1963. A systematic study of the Australian species of the genus Haemaphysalis Koch (Acarina: Ixodidae). Aust. J. Zool., 11: 35–80). Keirans and Clifford (1984, op. cit. under H. bartelsi) state that the USNTC “probably” contains paratypes of this species.

Known stages: male, female, larva

Zoogeographic Region: Australasian

Ecoregion: eastern Australia temperate forests

Hosts: Diprotodontia: Phalangeridae (ANL)

Artiodactyla: Bovidae; Perissodactyla: Equidae; Aegotheliformes: Aegothelidae (A)

Aves (unknown orders) (stages unknown)

Human infestation: no

Remarks: Kolonin (2009) states that the immature stages of H. bremneri have not been described, but there is a description of the larva in Roberts (1969). The nymph is undescribed, but the holotype of H. bremneri is from a nymph collected from Phalangeridae that molted into a male (Roberts 1963). Hoogstraal and Wassef (1973) and Hoogstraal and Kim (1985) state, without elaboration, that this tick may feed on various birds.

References

Heath, A.C.G. 1986. Aspects of the biology, seasonality and host associations of Haemaphysalis bancrofti, H. humerosa, H. bremneri and Ixodes tasmani (Acari: Ixodidae). In C. Cremin, C. Dobson & D.E. Moorhouse (editors), Parasite lives. University of Queensland Press, St Lucia, Australia, pp. 179–188.

Hoogstraal, H. & Kim, K.C. 1985. Tick and mammal coevolution, with emphasis on Haemaphysalis. In K.C. Kim (editor), Coevolution of parasitic arthropods and mammals. John Wiley & Sons, New York, pp. 505–568.

Hoogstraal, H. & Wassef, H.Y. 1973. The Haemaphysalis ticks (Ixodoidea: Ixodidae) of birds. 3.H. (Ornithophysalis) subgen. n.: definition, species, hosts and distribution in the Oriental, Palearctic, Malagasy, and Ethiopian Faunal Regions. J. Parasitol., 59: 1099–1117.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Roberts, F.H.S. 1963. A systematic study of the Australian species of the genus Haemaphysalis Koch (Acarina: Ixodidae). Aust. J. Zool., 11: 35–80.

Roberts, F.H.S. 1969, The larvae of Australian Ixodidae (Acarina: Ixodoidea). J. Aust. Entomol. Soc., 8: 37–78.

19 – H. calcarata Neumann, 1902 (Arch. Parasitol., 6: 109–128)

Type depository: BMNH (syntypes) (Keirans and Hillyard 2001, op. cit. under H. aciculifer)

Known stages: male, female, nymph

Zoogeographic Region: Afrotropical

Ecoregions: tropical and subtropical grasslands, savannas and shrublands

Hosts: usual hosts for adult ticks are Rodentia: Sciuridae.

Rodentia: Sciuridae (AN)

Carnivora: Felidae, Herpestidae (A)

Lagomorpha: Leporidae (N)

Rodentia: Muridae (stage unknown)

Human infestation: no

Remarks: Kolonin (2009) recognizes only Felidae as hosts for adult H. calcarata – not Sciuridae or Herpestidae. However, we consider records of adults of this tick on Herpestidae in Iori et al. (1996) to be valid.

References

Hoogstraal, H. 1955. Notes on African Haemaphysalis ticks. II. The ground-squirrel parasites, H. calcarata Neumann, 1902, and H. houyi Nuttall and Warburton, 1915 (Ixodoidea, Ixodidae). J. Parasitol., 41: 361–373.

Iori, A., Lanfrachi, P. & Manilla, G. 1996. Contribution to the knowledge of Ixodidae ticks of wild mammals of Somalia. Parassitologia, 38: 571–573.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Santos Dias, J.A.T. 1955. Contribuição para o conhecimento de fauna ixodológica do sudoeste africano. An. Inst. Med. Trop., 12: 75–100.

Theiler, G. 1962. The Ixodoidea parasites of vertebrates in Africa south of the Sahara (Ethiopian Region). Report to the Director of Veterinary Services, Onderstepoort, South Africa, Project S.9958, 260 pp.

20 – H. calva Nuttall & Warburton, 1915 (Ticks. A monograph of the Ixodoidea. Part III. The genus Haemaphysalis. Cambridge University Press, London, pp. 349–550)

Type depository: BMNH (lectotype, paralectotypes) (Keirans and Hillyard 2001, op. cit. under H. aciculifer) as Haemaphysalis calvus

Known stages: male, female, nymph

Zoogeographic Region: Oriental

Ecoregions: Sumatra peat swamp forests; Borneo rain forests

Hosts: usual hosts for adult ticks are Artiodactyla: Cervidae.

Artiodactyla: Bovidae, Cervidae, Suidae; Carnivora: Viverridae, Ursidae (A)

Human infestation: no

Remarks: the nymph of H. calva is only known from a nymphal pelt (Hoogstraal and Wassef 1981).

Reference

Hoogstraal, H. & Wassef, H.Y. 1981. Haemaphysalis (Garnhamphysalis) subgen. nov. (Acarina: Ixodidae): candidate tick vectors of hematozoa in the Oriental Region. Parasitol. Topics Spec. Publ., pp. 117–124.

21 – H. campanulata Warburton, 1908 (Proc. Cambr. Phil. Soc., 14: 508–519)

Type depository: BMNH (lectotype, paralectotypes) (Keirans and Hillyard 2001, op. cit. under H. aciculifer)

Known stages: male, female, nymph, larva

Zoogeographic Regions: Oriental, Palearctic

Ecoregions: temperate broadleaf and mixed forests

Hosts: usual hosts for larvae, nymphs and adult ticks are Carnivora: Canidae.

Carnivora: Canidae (ANL)

Mammalia (several orders) (A)

Rodentia: Muridae (NL)

Human infestation: yes (Yamaguti et al. 1971)

Remarks: Camicas et al. (1998) list the nymph of H. campanulata as undescribed, but it was described by Yamaguti et al. (1971). There is a record of this species’ introduction into the Australasian Region, but it failed to become established there (Roberts 1963). Noh (1972) records males, females and nymphs of H. campanulata on Galliformes: Phasianidae, while Phan Trong (1977) reports adults of this tick from Cuculiformes and Passeriformes, and Tsai et al. (2012) record one adult of H. campanulata from an undetermined bird, but we feel that these records require confirmation and they have not been included in our list of hosts for this species.

References

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. 1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.

Chen, Z., Yang, X., Bu, F., Yang, X., Yang, X. & Liu, J. 2010. Ticks (Acari: Ixodoidea: Argasidae, Ixodidae) of China. Exp. Appl. Acarol., 51: 393–404.

Hoogstraal, H. & Kim, K.C. 1985. Tick and mammal coevolution, with emphasis on Haemaphysalis. In K.C. Kim (editor), Coevolution of parasitic arthropods and mammals. John Wiley & Sons, New York, pp. 505–568.

Keirans, J.E. 1985. George Henry Falkiner Nuttall and the Nuttall tick catalogue. U. S. Dept. Agric., Agric. Res. Ser. Misc. Pub. (1438), 1785 pp.

Kitaoka, S. 1985. Keys to the species in immature stages of the Japanese Haemaphysalis ticks (Ixodidae). Bull. Natl. Inst. Anim. Health Q. (88): 49–63. In Japanese.

Noh, Y.T. 1972. Faunal studies on the ticks in Korea. In Floral studies on some taxa of plants and faunal studies on some taxa of animals in Korea. Ministry of Science and Technology, pp. 141–160. In Korean.

Nuttall, G.H.F. & Warburton, C. 1915. Ticks. A monograph of the Ixodoidea. Part III. The genus Haemaphysalis. Cambridge University Press, London, pp. 349–550.

Phan Trong, C. 1977. Ve bet va con trung ky sinh o Viet Nam. Tap 1. Ve (Ixodoidea), mo ta va phan loai. Ha Noi: Khoa hoc va ky thuat, 489 pp. In Vietnamese.

Roberts, F.H.S. 1963. A systematic study of the Australian species of the genus Haemaphysalis Koch (Acarina: Ixodidae). Aust. J. Zool., 11: 35–80.

Tsai, Y.-L., Shyu, C.-L., Yao, C.-T. & Lin, J.A. 2012. The ixodid ticks collected from dogs and other animals in Taiwan and Kinmen Island. Int. J. Acarol., 38: 110–115.

Yamaguti, N., Tipton, V.J., Keegan, H.L. & Toshioka, S. 1971. Ticks of Japan, Korea, and the Ryukyu Islands. Brigham Young Univ. Sci. Bull. Biol. Ser., 15 (1), 226 pp.

22 – H. canestrinii (Supino, 1897) (Atti Soc. Veneto-Trentina Sci. Nat. Residente Padova, Ser. 2, 3: 230–238)

Type depository: GM (lectotype, paralectotypes) (Hoogstraal and Trapido 1966, op. cit. under H. asiatica) as Opisthodon canestrinii

Known stages: male, female, nymph

Zoogeographic Regions: Oriental, Palearctic

Ecoregions: tropical and subtropical forests

Hosts: usual hosts for adult ticks are Carnivora: Canidae, Felidae and Viverridae. Aves are considered exceptional hosts for adults of this tick.

Carnivora: Viverridae (AN)

Carnivora (several families); Lagomorpha: Leporidae; Galliformes: Phasianidae (A)

Rodentia: Sciuridae, Muridae; Scandentia: Tupaiidae (N)

Human infestation: no

Remarks: data published prior to Hoogstraal (1971) have been ignored because of uncertainties concerning the identification of H. canestrinii. Camicas et al. (1998) treat H. canestrinii as an exclusively Oriental species, but several records in Teng and Jiang (1991) are from the Palearctic Region. Hoogstraal (1971) records this species on humans, but the specimens were not actually feeding, and we have therefore not included humans as hosts of H. canestrinii. Kolonin (2009) ignores Aves as hosts of this species, but we regard the odd record from Aves in Hoogstraal (1971) as sound. Kolonin (2009) appears to believe that the larvae of H. canestrinii feed on various host species, but we have been unable to confirm this.

References

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. 1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.

Hoogstraal, H. 1971. Identity, hosts, and distribution of Haemaphysalis (Rhipistoma) canestrinii (Supino) (resurrected), the postulated Asian progenitor of the African leachi complex (Ixodoidea: Ixodidae). J. Parasitol., 57: 161–172.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Tanskul, P., Stark, H.E. & Inlao, I. 1983. A checklist of ticks of Thailand (Acari: Metastigmata: Ixodoidea). J. Med. Entomol., 20: 330–341.

Teng, K.-F. & Jiang, Z.-J. 1991. Economic insect fauna of China. Fasc. 39, Acari: Ixodidae. Science Press, Beijing, 355 pp. In Chinese.

23 – H. capricornis Hoogstraal, 1966 (J. Parasitol., 52: 783–786)

Type depositories: BMNH (holotype, paratypes), USNTC (paratypes) (Keirans and Hillyard 2001, op. cit. under H. aciculifer), originally identified as H. hystricis, as stated in Hoogstraal, H. (1966. Studies on Southeast Asian Haemaphysalis ticks (Ixodoidea: Ixodidae). H. (H.) capricornis sp. n., the large Malayan serow haemaphysalid from southwestern Thailand. J. Parasitol., 52: 783–786)

Known stages: male

Zoogeographic Region: Oriental

Ecoregion: Tenasserim-South Thailand semi-evergreen rain forests

Hosts: Artiodactyla: Bovidae (A)

Human infestation: no

Remarks: Tanskul et al. (1983) list Muridae as hosts for the adults of H. capricornis, but this record is not included in Tanskul and Inlao (1989) (both papers refer to ticks in Thailand), and we have therefore omitted murids from our host list for this tick. Mihalca et al. (2011) regard H. capricornis as endangered.

References

Hoogstraal, H. 1966. Studies on Southeast Asian Haemaphysalis ticks (Ixodoidea: Ixodidae). H. (H.) capricornis sp. n., the large Malayan serow haemaphysalid from southwestern Thailand. J. Parasitol., 52: 783–786.

Mihalca, A.D., Gherman, C.M. & Cozma, V. 2011. Coendangered hard-ticks: threatened or threatening? Parasit. Vectors, 4 (71), 7 pp.

Tanskul, P. & Inlao, I. 1989. Keys to the adult ticks of Haemaphysalis Koch, 1844, in Thailand with notes on changes in taxonomy (Acari: Ixodoidea: Ixodidae). J. Med. Entomol., 26: 573–601.

Tanskul, P., Stark, H.E. & Inlao, I. 1983. A checklist of ticks of Thailand (Acari: Metastigmata: Ixodoidea). J. Med. Entomol., 20: 330–341.

24 – H. caucasica Olenev, 1928 (Dokl. Akad. Nauk USSR, Ser. A (2): 29–34. In Russian)

Type depository: ZIAC (lectotype, paralectotype) (Filippova, N.A. 2008. Type specimens of argasid and ixodid ticks (Ixodoidea: Argasidae, Ixodidae) in the collection of the Zoological Institute, Russian Academy of Sciences (St. Petersburg). Entomol. Rev., 88: 1002–1011)

Known stages: male, female, nymph, larva

Zoogeographic Region: Palearctic

Ecoregions: desert and xeric shrublands

Hosts: usual hosts for larvae, nymphs and adults are Lagomorpha: Leporidae.

Lagomorpha: Leporidae (ANL)

Artiodactyla: Bovidae; Carnivora: Canidae, Mustelidae, Ursidae (A)

Erinaceomorpha: Erinaceidae; Rodentia: Muridae; Passeriformes (several families);

Coraciiformes: Upupidae; Galliformes: Phasianidae; Squamata: Lacertidae, Scincidae (NL)

Human infestation: yes (Filippova 1997)

Remarks: Camicas et al. (1998) consider pholeophilic (burrowing) mammals to be the sole hosts of this tick species, but we believe that Aves are important hosts for the larvae and nymphs of H. caucasica, as stated in Filippova (1997). Durden and Keirans (1996) list H. caucasica as an endangered species.

References

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. 1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.

Durden, L.A. & Keirans, J.E. 1996. Host-parasite coextinction and the plight of tick conservation. Am. Entomol., 42: 87–91.

Filippova, N.A. 1997. Ixodid ticks of subfamily Amblyomminae. Fauna of Russia and neighbouring countries, 4 (5), Nauka, St. Petersburg, 436 pp.

25 – H. celebensis Hoogstraal, Trapido & Kohls, 1965 (J. Parasitol., 51: 1001–1003)

Type depository: BMNH (holotype) (Keirans and Hillyard 2001, op. cit. under H. aciculifer). Originally named H. hystricis in Nuttall, G.H.F. & Warburton, C. (1915. Ticks. A monograph of the Ixodoidea. Part III. The genus Haemaphysalis. Cambridge Univ. Press, London, pp. 349–550), as stated in Hoogstraal, H., Trapido, H. & Kohls, G.M. (1965. Studies on Southeast Asian Haemaphysalis ticks (Ixodoidea, Ixodidae). H. (Kaiseriana) celebensis sp. n., from a wild boar in Celebes. J. Parasitol., 51, 1001–1003)

Known stages: male, female

Zoogeographic Region: Australasian

Ecoregion: Sulawesi montane rain forests

Hosts: usual hosts for adult ticks are Artiodactyla: Suidae.

Artiodactyla: Bovidae, Cervidae, Suidae; Perissodactyla: Equidae (A)

Rodentia: Muridae (NL)

Human infestation: yes (Durden et al. 2008)

Remarks: the larva and nymph of H. celebensis have not been described, but we consider records of the immature stages of this tick in Durden et al. (2008) to be provisionally valid. Kolonin (2009) ignores Equidae as hosts for H. celebensis, but we accept the record from this type of host in Durden et al. (2008).

References

Durden, L.A., Merker, S. & Beati, L. 2008. The tick fauna of Sulawesi, Indonesia (Acari: Ixodoidea: Argasidae and Ixodidae). Exp. App. Acarol., 45: 85–110.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

26 – H. chordeilis (Packard, 1869) (First Annual Report of the Trustees of the Peabody Academy of Sciences, Appendix, pp. 52–69)

Type depository: MCZ (holotype). Guglielmone, A.A., Estrada-Peña, A., Keirans, J.E. & Robbins, R.G. (2003. Ticks (Acari: Ixodida) of the Neotropical Zoogeographic Region. Special Publication of the International Consortium on Ticks and Tick-borne Diseases-2, Atalanta, Houten, The Netherlands, 173 pp.) state that the types were housed in MPAS as Ixodes chordeilis but have apparently been lost. However, Cooley, R.A. (1946. The genera Boophilus, Rhipicephalus, and Haemaphysalis (Ixodoidea) of the New World. Natl. Inst. Health Bull. (187), 54 pp.) states that “types” are in MCZ, as Ixodes chordeilis. The MCZ online zoological collections database (http://mczbase.mcz.harvard.edu/ accessed April 4, 2013) indicates that the holotype of H. chordeilis is held there.

Known stages: male, female, nymph, larva

Zoogeographic Region: Nearctic

Ecoregions: several different Nearctic ecoregions

Hosts: Mammalia are considered exceptional hosts for this tick.

Galliformes: Phasianidae; Passeriformes: Icteridae (ANL)

Anseriformes: Anatidae; Caprimulgiformes: Caprimulgidae; Artiodactyla: Bovidae; Perissodactyla: Equidae; Rodentia: Sciuridae (A)

Passeriformes: Emberizidae (NL)

Falconiformes: Accipitridae; Passeriformes: Mimidae (N)

Passeriformes: Corvidae, Troglodytidae (L)

Rodentia: Geomyidae (stage unknown)

Human infestation: yes (Keirans 1985)

Remarks: Camicas et al. (1998) state that H. chordeilis is present in the Nearctic and Neotropical Regions, but there are no bona fide Neotropical records of this species, as discussed in Guglielmone et al. (2003). Cooley (1946) cites Hewitt (1915, who uses the name H. cinnabarina) concerning Leporidae as hosts of H. chordeilis, while Allan (2001) lists the domestic cat as a host without supplying details for this record. Becklund (1964) presents a table with the caption “parasites of dog and cats” in which H. chordeilis is included, but without specifying whether the tick was a parasite of Canidae, Felidae or both. None of the latter four authors mention the developmental stage of ticks found on hosts. These doubtful records have been excluded from our list of hosts of H. chordeilis. Infestation of mammals with H. chordeilis does not appear to be a common event. See also H. cinnabarina.

References

Allan, S.A. 2001. Ticks (Class Arachnida: Order Acarina). In W.S. Samuel, M.J. Pybus & A.A. Kocan (editors), Parasitic Diseases of Wild Animals, 2^nd edition. Iowa State University Press, Ames, pp. 72–106.

Becklund, W.W. 1964. Revised check list of internal and external parasites of domestic animals in the United States and possessions and in Canada. Am. J. Vet. Res., 25: 1380–1416.

Bequaert, J.C. 1945. The ticks, or Ixodoidea, of the northeastern United States and eastern Canada. Entomol. Am., 25: 73–232.

Bishopp, F.C. & Trembley, H.L. 1945. Distribution and hosts of certain North American ticks. J. Parasitol., 31: 1–54.

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. 1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.

Cooley, R.A. 1946. The genera Boophilus, Rhipicephalus, and Haemaphysalis (Ixodidae) of the New World. Natl. Inst. Health Bull. (187), 54 pp.

Gregson, J.D. 1956. The Ixodoidea of Canada. Can. Dept. Agric. Sci. Serv. Entomol. Div. Publ. (930), 92 pp.

Guglielmone, A.A., Estrada-Peña, A., Keirans, J.E. & Robbins, R.G. 2003. Ticks (Acari: Ixodida) of the Neotropical Zoogeographic Region. Special Publication of the International Consortium on Ticks and Tick-borne Diseases-2, Atalanta, Houten, The Netherlands, 173 pp.

Hewitt, C.G. 1915. A contribution to a knowledge of Canadian ticks. Trans. R. Soc. Can. Ser. 3, 9: 225–239.

Keirans, J.E. 1967. Some avian ectoparasites in New England. Entomol. News, 78: 40–42.

Keirans, J.E. 1985. George Henry Falkiner Nuttall and the Nuttall tick catalogue. U. S. Dept. Agric., Agric. Res. Ser. Misc. Pub. (1438), 1785 pp.

Wilson, N. & Kale, H.W. 1972. Ticks collected from Indian River County, Florida (Acari: Metastigmata: Ixodidae). Florida Entomol., 55: 53–57.

27 – H. cinnabarina Koch, 1844 (Arch. Naturgesch., 10: 217–239)

This species is not included in Kolonin (2009, op. cit. under H. anomaloceraea). See also remarks below.

Type depository: ZMB (holotype) (Moritz, M. & Fischer, S.C. 1981. Die Typen der Arachniden-Sammlung des Zoologischen Museums Berlin. Mitt. Zool. Mus. Berlin, 57: 341–364)

Known stages: female

Zoogeographic Region: Neotropical

Ecoregion: the only known records mention Brazil as locality, without further geographical information

Hosts: unknown

Human infestation: no

Remarks: this tick is known from two specimens collected almost 170 years ago as H. cinnabarina and its synonym H. sanguinolenta Koch, 1844. Although Hoogstraal (1973) defended the Brazilian origin of H. cinnabarina, Guglielmone et al. (2003) did not include it as a Neotropical taxon. Their opinion was based on the statement by Keirans and Restifo (1993) that most investigators consider this name identical to H. punctata, and Kolonin (2009) also maintains this view. Cooley (1946) had previously believed that H. cinnabarina was partly a synonym of H. chordeilis. Recently, Barros-Battesti et al. (2008) examined the type and concurred with the opinion of Hoogstraal (1973). Consequently, we now regard H. cinnabarina as a tick exclusive to the Neotropical Zoogeographic Region. Camicas et al. (1998) list ungulates as hosts of this tick species, but we have found no information to support this statement. There is a record from Rodentia in Suriname (Neotropical Region) by Reyne (1923), but it requires confirmation and we have excluded it from our list of bona fide hosts of H. cinnabarina.

References

Barros-Battesti, D.M., Onofrio, V.C., Arzua, M. & Labruna, M.B. 2008. Comments on the validity of Haempahysalis cinnabarina Koch, 1844 (Acari: Ixodidae), a taxon known solely by the type specimens from northern Brazil. Rev. Bras. Parasitol. Vet., 17 (1): 53–55.

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. 1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.

Cooley, R.A. 1946. The genera Boophilus, Rhipicephalus, and Haemaphysalis (Ixodidae) of the New World. Natl. Inst. Health Bull. (187), 54 pp.

Guglielmone, A.A., Estrada-Peña, A., Keirans, J.E. & Robbins, R.G. 2003. Ticks (Acari: Ixodida) of the Neotropical Zoogeographic Region. Special Publication of the International Consortium on Ticks and Tick-borne Diseases-2, Atalanta, Houten, The Netherlands, 173 pp.

Hoogstraal, H. 1973. Redescription of the type material of Haemaphysalis (Aboimisalis) cinnabarina (revalidated) and its junior synonym H. (A.) sanguinolenta described by Koch in 1844 from Brazil (Ixodoidea: Ixodidae). J. Parasitol., 59: 379–383.

Keirans, J.E. & Restifo, R.A. 1993. Haemaphysalis juxtakochi Cooley (Acari: Ixodidae), a Neotropical tick species, found in Ohio. J. Med. Entomol., 30: 1074–1075.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Reyne, A. 1923. Verslag van den entomoloog. Versl. Dep. Landbouw. Nijver. Handel Suriname (1922), pp. 32–39.

28 – H. colasbelcouri (Santos Dias, 1958) (Mem. Estud. Mus. Zool. Univ. Coimbra (249), 9 pp.)

Barker, S.C. & Murrell, A. (2008. Systematics and evolution of ticks with a list of valid genus and species names. In A.S. Bowman & P.A. Nuttall (editors), Ticks: biology, disease and control. Cambridge University Press, Cambridge, pp. 1–39) include as valid Haemaphysalis vietnamensis Hoogstraal & Wilson, 1966 and H. colasbelcouri, but the former species has since been treated as a synonym of the latter (Guglielmone, A.A., Robbins, R.G., Apaneskevich, D.A., Petney, T.N., Estrada-Peña, A. & Horak, I.G., 2009. Comments on controversial tick (Acari: Ixodida) species names and species described or resurrected from 2003 to 2008. Exp. Appl. Acarol., 48: 311–327). Kolonin (2009, op. cit. under H. anomaloceraea) includes this name as? H. colasbelcouri under the name I. vietnamensis without further explanation. Chen, Z., Yang, X., Bu, F., Yang, X., Yang, X. & Liu, J. (2010. Ticks (Acari: Ixodoidea: Argasidae, Ixodidae) of China. Exp. Appl. Acarol., 51: 393–404.) listed H. vietnamensis but not H. colasbelcouri for China, probably because they accepted Kolonin’s view (op. cit. above).

Type depository: IPP (holotype, paratypes) (Santos Dias, J.A.T. 1958. Notas ixodológicas. III. Sobre duas espécies do género Aponomma Neumann, 1899 da Região Oriental. Mem. Estud. Mus. Zool. Univ. Coimbra (249): 1–9) as Aponomma colasbelcouri

Known stages: male, female, nymph, larva

Zoogeographic Region: Oriental

Ecoregions: tropical and subtropical dry broadleaf forests

Hosts: Artiodactyla: Bovidae, Cervidae (A)

Human infestation: yes (Kolonin 1995 as H. vietnamensis, see above)

Remarks: Camicas et al. (1998) list the nymph and larva of this species as undescribed, but Ding and Ying (1996) described them under the name H. vietnamensis. The latter authors appear to have described these immature stages from laboratory-reared specimens, although we have only seen the abstract of their paper. Mihalca et al. (2011) regard this species as endangered under the name H. vietnamensis.

References

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. 1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.

Ding, X.-C. & Ying, P.-Y. 1996. Scanning electron microscopic observations of Haemaphysalis vietnamensis. Chin. J. Vet. Sci., 16: 486–490. In Chinese.

Kolonin, G.V. 1995. Review of the ixodid tick fauna (Acari: Ixodidae) of Vietnam. J. Med. Entomol., 32: 276–282.

Mihalca, A.D., Gherman, C.M. & Cozma, V. 2011. Coendangered hard-ticks: threatened or threatening? Parasit. Vectors, 4 (71), 7 pp.

29 – H. colesbergensis Apanaskevich & Horak, 2008 (J. Parasitol., 94: 594–607)

Kolonin (2009, op. cit. under H. anomaloceraea) does not include this species in his list of Ixodidae of the world.

Type depositories: USNTC (holotype, paratypes), OVI, ZIAC (paratypes) (Apanaskevich, D.A. & Horak, I.G. 2008. Two new species of African Haemaphysalis ticks (Acari: Ixodidae), carnivore parasites of the H. (Rhipistoma) leachi group. J. Parasitol., 94: 594–607). This tick was originally identified as H. leachi.

Known stages: male, female, nymph, larva

Zoogeographic Region: Afrotropical

Ecoregions: mediterranean woodlands and scrub; deserts and xeric shrublands

Hosts: Erinaceomorpha: Erinaceidae (AN)

Carnivora: Canidae, Felidae (A)

Human infestation: no

Remarks: the natural hosts for the larva of H. colesbergensis are unknown, and the larva and nymph were described from laboratory-reared material.

References

Apanaskevich, D.A. & Horak, I.G. 2008. Two new species of African Haemaphysalis ticks (Acari: Ixodidae), carnivore parasites of the H. (Rhipistoma) leachi group. J. Parasitol., 94: 594–607.

Horak, I.G., Welman, S., Hallam, S.L., Lutermann, H. & Mzilikazi, N. 2011. Ticks of four-toed elephant shrews and Southern African hedgehogs. Onderstepoort J. Vet. Res. 78, doi:10.4102/ojvr.v78i1.243

30 – H. concinna Koch, 1844 (Arch. Naturgesch., 10: 217–239)

Type depository: ZMB (syntypes) (Nuttall and Warburton 1915, op. cit. under H. celebensis), but not included in ZMB (Moritz and Fischer 1981, op. cit. under H. cinnabarina)

Known stages: male, female, nymph, larva

Zoogeographic Regions: Oriental, Palearctic

Ecoregions: several different ecoregions within the Palearctic and Oriental Zoogeographic Regions

Hosts: Squamata are considered exceptional hosts for this tick.

Mammalia (several orders); Galliformes: Phasianidae; Passeriformes (several families) (ANL)

Aves (several orders); Squamata: Colubridae, Lacertidae (NL)

Human infestation: yes (Hillyard 1996; Bursali et al. 2012 among others)

Remarks: recent records of H. concinna from China indicate that its distribution includes the Oriental Region (Chu et al. 2008; Bian et al. 2009). These records are not included in Chen et al. (2010), but we treat them as valid. Camicas et al. (1998) do not list Aves as significant hosts for larvae and nymphs of H. concinna, but Kislenko and Koneva (1965), Nosek (1971) and others found Aves to be important hosts of this species. Kolonin (2009) excludes Aves as hosts for adult H. concinna, and Squamata are also ignored as hosts of this tick. We accept the records of H. concinna adults on Aves in Kislenko and Koneva (1965), as well as the unusual records on Squamata in Nosek (1971) and Krčmar (2012).

References

Bian, Y., Yang, G., Wang, T., Yan, H., Wang, S., Sun, J. & Li, K. 2009. Haemaphysalis concinna (Acari: Ixodida): Persistent efficacy of doramectin in rabbits under laboratory conditions. Entomol. Res., 39: 119–122.

Bursali, A., Keskin, A. & Tekin, S. 2012. A review of the ticks (Acari: Ixodida) of Turkey: species diversity, hosts and geographical distribution. Exp. Appl. Acarol., 57: 91–104

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. 1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.

Chen, Z., Yang, X., Bu, F., Yang, X., Yang, X. & Liu, J. 2010. Ticks (Acari: Ixodoidea: Argasidae, Ixodidae) of China. Exp. Appl. Acarol., 51: 393–404.

Chu, C.-Y., Jiang, B.-G., Liu, W., Zhao, Q.-M., Wu, X.-M., Zhang, P.-H., Zhan, L., Yang, H. & Cao, W.-C. 2008. Presence of pathogenic Borrelia burgdorferi sensu lato in ticks and rodents in Zhejiang, south-east China. J. Med. Microbiol., 57: 980–985.

Emel’yanova, N.D. & Gordeeva, V.P. 1969. Data on the study of birds acting as hosts of ixodid ticks in East Manchurian mountain area spurs and in the Pre-Khanka Plain. In A. O. Cherepanov (editor), Migratory birds and their role in arboviral distribution. Sibirisk. Otd. Akad. Nauk SSSR, Biol. Inst., Akad. Nauk SSSR, Inst. Polio. Virus Entsef., Min. Zdravookhr. RSFSR, Omsk Inst. Prirod.-Ochag. Infekts., Novosibirsk, pp. 160–169. In Russian, NAMRU-3 translation 621.

Feider, Z. 1964. Les connaissances actuelles sur les Acariens de Roumanie (Ixodides, Gamasides et Trombiculides). Acarologia (Fasc. Hors Sér.): 262–274.

Filippova, N.A. 1997. Ixodid ticks of subfamily Amblyomminae. Fauna of Russia and neighbouring countries, 4 (5), Nauka, St. Petersburg, 436 pp. In Russian.

Hillyard, P.D. 1996. Ticks of North-West Europe. Keys and notes for identification of the species. Synopses of the British Fauna (New Series) (52), Field Studies Council, Shrewsbury, UK, 178 pp.

Kislenko, G.S. & Koneva, I.V. 1965. Infestations of birds by ixodid ticks in secondary forests in lower reaches of the Ussuri River. Mater. 4. Vses. Orinthol. Konf., Alma-Ata, Sep. 1965, pp. 160–163. In Russian, NAMRU-3 translation 408.

Krčmar, S. 2012. Hard ticks (Acari, Ixodidae) of Croatia. ZooKeys, 234: 19–57.

Lebedeva, N.N. & Korenberg, E.I. 1981. Distribution of Haemaphysalis concinna Koch in the Soviet Union and some general features of its ecology. Folia Parasitol., 28: 249–261.

Morel, P.-C. 2003. Les tiques d’Afrique et du Bassin méditerranéen (1965–1995). CIRAD- EMVT, 1342 pp.

Nosek, J. 1971. The ecology, bionomics and behaviour of Haemaphysalis (Haemaphysalis) concinna tick. Z. Parasitenkd., 36: 223–241.

Serdjukova, G.V. 1956. Ixodid ticks of the fauna of USSR. Opred. Faune SSSR. Zool. Inst. Akad. Nauk SSSR (64), 121 pp. In Russian.

Voloshina, I.V. 2010. Ticks (Ixodidae) of the Lazovsky Reserve. Proc. Sci. Conf devoted 75 Anniv. Lazovsky Reserve, Vladivostok, pp. 59–65. In Russian.

31 – H. cooleyi Bedford, 1929 (15^th Annual Report of the Director of Veterinary Services, Department of Agriculture, Union of South Africa, pp. 493–499)

See remarks below.

Type depository: OVI (lectotype, paralectotypes) (Hoogstraal, H. & Wassef, H.Y. 1981. Notes on African Haemaphysalis ticks. XIII. Identity of H. (Rhipistoma) cooleyi, a parasite of the rock hyrax in South Africa (Acarina: Ixodidae). Onderstepoort J. Vet. Res., 48: 135–140)

Known stages: male, female, nymph

Zoogeographic Region: Afrotropical

Ecoregions: rocky outcrops and mountains in subtropical savannas and shrublands

Hosts: Hyracoidea: Procaviidae (AN)

Human infestation: no

Remarks: Bedford (1929) includes nymphs of H. cooleyi from Rodentia: Pedetidae in his description of this species. However, as reported in Hoogstraal (1956), these nymphs were in fact a male and a female that had at first been classified as H. numidiana Neumann, 1905 (= H. erinacei). Subsequently, Hoogstraal (1972) decided that these specimens belonged to a new species (H. pedetes). Descriptions and illustrations of the male, female and nymph of H. cooleyi are included in Hoogstraal & Wassef (1981).

References

Bedford, G.A.H. 1929. Notes on some South African ticks, with descriptions of three new species. 15^th Annual Report of the Director of Veterinary Services, Department of Agriculture, Union of South Africa, pp. 493–499.

Hoogstraal, H. 1956. Notes on African Haemaphysalis ticks. III. The hyrax parasites, H. bequaerti sp. nov., H. orientalis N. and W., 1915 (new combination), and H. cooleyi Bedford, 1929 (Ixodoidea, Ixodidae). J. Parasitol., 42: 156–172.

Hoogstraal, H. 1972. Notes on African Haemaphysalis ticks. IX. H. (Rhipistoma) pedetes sp. n., a relict parasite of the springhare (Rodentia: Pedetidae) in Transvaal (Ixodoidea: Ixodidae). J. Parasitol., 58: 979–983.

Hoogstraal, H. & Wassef, H.Y. 1981. Notes on African Haemaphysalis ticks. XIII. Identity of H. (Rhipistoma) cooleyi, a parasite of the rock hyrax in South Africa (Acarina: Ixodidae). Onderstepoort J. Vet. Res., 48: 135–140.

Walker, J.B. 1991. A review of the ixodid ticks (Acari, Ixodidae) occurring in southern Africa. Onderstepoort J. Vet. Res., 58: 81–105.

32 – H. cornigera Neumann, 1897 (Mém. Soc. Zool. Fr., 10: 324–420)

See H. taiwana and remarks below.

Type depositories: MNHN (lectotype), BMNH (paralectotype) (Keirans & Hillyard, 2001, op. cit. under H. aciculifer)

Known stages: male, female, nymph, larva

Zoogeographic Regions: Australasian, Oriental, Palearctic

Ecoregions: tropical and subtropical moist broadleaf forests

Hosts: Aves are considered exceptional hosts for adults of this tick.

Carnivora: Viverridae (ANL)

Artiodactyla: Bovidae, Cervidae; Rodentia: Muridae (AN)

Artiodactyla: Suidae; Carnivora: Canidae, Felidae, Mustelidae; Rodentia: Hystricidae; Passeriformes: Timaliidae, Turdidae; Galliformes: Phasianidae (A)

Scandentia: Tupaiidae (N)

Rodentia: Sciuridae; Soricomorpha: Soricidae (N?)

Human infestation: yes (Durden et al. 2008)

Remarks: there is considerable confusion concerning H. cornigera and H. ias (see H. ias for further details) and contradictory information about H. cornigera itself. The larva and nymph of H. cornigera are listed as undescribed in Camicas et al. (1998), while Anastos (1950) initially expressed doubts about the Oudemans’ (1927) descriptions of the larva and nymph based on specimens from Ambon Island (Australasian), only to later accept this locality. Durden et al. (2008) also found H. cornigera in Sulawesi (Australasian). The descriptions of the sub-adult stages of H. cornigera by Oudemans (1927) are sufficiently detailed and we have found no direct criticism of his work. Consequently, we regard his descriptions as provisionally valid. Camicas et al. (1998) state that H. cornigera occurs in the Oriental and Palearctic Zoogeographic Regions. However, we were unable to find bona fide Palearctic localities for this species prior to Yamauchi et al. (2009), who recorded its presence in Honshu (Japan). Because Ambon Island and Sulawesi lie within the Australasian Zoogeographic Region, the distribution of H. cornigera includes this region in addition to the Oriental and Palearctic Regions listed by Camicas et al. (1998). Tanskul et al. (1983) present information on the hosts of H. cornigera in Thailand, but Tanskul & Inlao (1989) later state that the ticks identified as H. cornigera in Thailand were in fact H. shimoga. Sreenivasan et al. (1972) generalized that Soricomorpha and Rodentia: Muridae and Sciuridae were hosts for nymphs of H. cornigera, but it is unclear from their publication whether all these hosts were infested. Kolonin (2009) lists Muridae as hosts for the immature stages of H. cornigera, probably following Hoogstraal et al. (1972), who refer to this tick as H. cornigera cornigera, and state that nymphs and larvae feed on rodents, but present information for three nymphs from vegetation. We treat the record of Oudemans (1927) of larvae and nymphs on Viverridae, the record of Warburton (1926) of nymphs on Tupaiidae, and the record of Munaf (1978) of a nymph on Muridae as provisionally valid. Kolonin (2009) does not include Aves as hosts for adult H. cornigera, but we consider the few records from Aves in Anastos (1950) to be provisionally valid. Clearly, the literature on H. cornigera contains a number of conflicting statements.

References

Anastos, G. 1950. The scutate ticks, or Ixodidae, of Indonesia. Entomol. Am., 30: 1–144.

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. 1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.

Chen, Z., Yang, X., Bu, F., Yang, X., Yang, X. & Liu, J. 2010. Ticks (Acari: Ixodoidea: Argasidae, Ixodidae) of China. Exp. Appl. Acarol., 51: 393–404.

Durden, L.A., Merker, S. & Beati, L. 2008. The tick fauna of Sulawesi, Indonesia (Acari: Ixodoidea: Argasidae and Ixodidae). Exp. App. Acarol., 45: 85–110.

Hoogstraal, H., Lim, B.L., Nadchatram, M. & Anastos, G. 1972. The Gunong Benom Expedition 1967. 8. Ticks (Ixodidae) of Gunong Benom and their altitudinal distribution, hosts and medical relationships. Bull. Br. Mus. (Nat. Hist.) Zool., 23: 167–186.

Keirans, J.E. 1985. George Henry Falkiner Nuttall and the Nuttall tick catalogue. U. S. Dept. Agric., Agric. Res. Ser. Misc. Pub. (1438), 1785 pp.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Munaf, H.B. 1978. Tick fauna of Baluran Wildlife Reserve, Indonesia. Hemera Zoa, 70: 37–44.

Oudemans, A.C. 1927. X. Acari uit Ambon. Zool. Meded. Rijks Mus. Nat. Hist., 10: 185–237.

Phan Trong, C. 1977. Ve bet va con trung ky sinh o Viet Nam. Tap 1. Ve (Ixodoidea), mo ta va phan loai. Ha Noi: Khoa hoc va ky thuat, 489 pp. In Vietnamese.

Sreenivasan, M.A., Rajagopalan, P.K. & Bhat, H.R. 1972. Isolation of Kyasanur Forest disease virus from ixodid ticks collected between 1965 and 1972. In G.P. Channabasavanna & C.A. Viraktamath (editors), Progress in acarology, volume I. Oxford & IBH Publishing Company, New Dehli, pp. 37–44.

Tanskul, P. & Inlao, I. 1989. Keys to the adult ticks of Haemaphysalis Koch, 1844, in Thailand with notes on changes in taxonomy (Acari: Ixodoidea: Ixodidae). J. Med. Entomol., 26: 573–601.

Tanskul, P., Stark, H.E. & Inlao, I. 1983. A checklist of ticks of Thailand (Acari: Metastigmata: Ixodoidea). J. Med. Entomol., 20: 330–341.

Warburton, C. 1926. Ixodidae from Sumatra. Treubia, 8: 280.

Yamauchi, T., Tabara, K., Kanamori, H., Kawabata, H., Arai, S., Katayama, T., Fujita, H., Yano, Y., Takada, N. & Itagaki, A. 2009. Tick fauna associated with sika deer density in the Shimane Peninsula, Honshu. Japan. Med. Entomol Zool., 60: 297–304.

33 – H. cornupunctata Hoogstraal & Varma, 1962 (J. Parasitol., 48: 185–194)

Type depositories: USNTC (holotype, paratypes), BMNH, HH, VRC, ZSI (paratypes) (Keirans & Hillyard, 2001, op. cit. under H. aciculifer)

Known stages: male, female, nymph, larva

Zoogeographic Regions: Oriental, Palearctic

Ecoregions: montane grasslands and shrublands

Hosts: usual hosts for larvae, nymphs and adults are Artiodactyla: Bovidae.

Artiodactyla: Bovidae; Perissodactyla: Equidae (ANL)

Rodentia: Muridae (N)

Lagomorpha: Ochotonidae (stage unknown)

Human infestation: no

Remarks: Camicas et al. (1998) consider this species to be exclusively Oriental, but Hoogstraal and Kim (1985) include Afghanistan (Palearctic) within its range. We were unable to confirm the record of H. cornupunctata on Cervidae in Ghosh et al. (2007). Rao et al. (1973) state that “mouse hare” is a host of H. cornupunctata but do not identify the tick stage(s) collected. Kolonin (2009) does not recognize hosts of H. cornupunctata other than “livestock.”

References

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. 1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.

Dhanda, V. & Kulkarni, S.M. 1969. Immature stages of Haemaphysalis cornupunctata Hoogstraal & Varma, 1962 (Acarina: Ixodidae) with new host and locality records, and notes on its ecology. Oriental Insects, 3: 15–21.

Ghosh, S., Bansal, G.C., Gupta, S.C., Ray, D., Khan, M.Q., Irshad, H., Shahiduzzaman, Md., Seitzer, U. & Ahmed, J.S. 2007. Status of tick distribution in Bangladesh, India and Pakistan. Parasitol. Res., 101 (Suppl. 2): 207–216.

Hoogstraal, H. & Kim, K.C. 1985. Tick and mammal coevolution, with emphasis on Haemaphysalis. In K.C. Kim (editor), Coevolution of parasitic arthropods and mammals. John Wiley & Sons, New York, pp. 505–568.

Hoogstraal, H. & Varma, M.G.R. 1962. Haemaphysalis cornupunctata sp. n. and H. kashmirensis sp. n. from Kashmir, with notes on H. sundrai Sharif and H. sewelli Sharif of India and Pakistan (Ixodoidea: Ixodidae). J. Parasitol., 48: 185–194.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Rao, T.R., Dhanda, V., Bhat, H.R. & Kulkarni, S.M. 1973. A survey of haematophagous arthropods in western Himalayas, Sikkim and Hill Districts of West Bengal. A general account. Ind. J. Med. Res., 61: 1421–1461.

Sharma, B.D. 1993. Medical & veterinary arthropod-disease ecology. Ashish Publishing House, New Delhi, 335 pp.

34 – H. cuspidata Warburton, 1910 (Parasitology, 3: 395–407)

Type depositories: BMNH (lectotype, paralectotypes), ZMB (paratype) [sic] (Keirans & Hillyard, 2001, op. cit. under H. aciculifer)

Known stages: male, female, nymph, larva

Zoogeographic Region: Oriental

Ecoregions: Sri Lanka and southwestern Ghats montane rain forests

Hosts: Artiodactyla: Tragulidae; Carnivora: Viverridae; Cuculiformes: Cuculidae (ANL) Carnivora: Felidae, Herpestidae; Lagomorpha: Leporidae (A)

Artiodactyla: Bovidae; Primates: Cercopithecidae (NL)

Rodentia: Hystricidae; Coraciiformes: Bucerotidae; Galliformes: Phasianidae (N)

Rodentia: Muridae, Sciuridae; Soricomorpha: Soricidae; Passeriformes: Sylviidae (L)

Carnivora: Canidae (stage unknown)

Human infestation: no

Remarks: Kolonin (2009) does not include Aves as hosts of adults of H. cuspidata, but we regard the records from Cuculidae in Rajagopalan (1972) as sound. Kolonin (2009) also lists Cervidae as hosts for all parasitic stages of H. cuspidata, but we were unable to confirm this from the reference provided by this author and have therefore excluded Cervidae from our host list above.

References

Bhat, H.R. & Sreenivasan, M.A. 1981. Further records of the ticks of some reptilian and mammalian hosts in the Kyasanur Forest disease area, Karnataka, India. Ind. J. Parasitol., 5: 207–210.

Boshell, J. & Rajagopalan, P.K. 1968. Observations on the experimental exposure of monkeys, rodents and shrews to infestation of ticks in forest in Kyasanur Forest disease area. Ind. J. Med. Res., 56 (4) (Suppl.): 573–588.

Grokhovskaya, I.M. & Nguen, H.H. 1968. Contribution to the study of ixodid ticks (Ixodidae) in Vietnam. Med. Parazit., 37: 710–715. In Russian, NAMRU-3 translation 401.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Nuttall, G.H.F. & Warburton, C. 1915. Ticks. A monograph of the Ixodoidea. Part III. The genus Haemaphysalis. Cambridge University Press, London, pp. 349–550.

Rajagopalan, P.K. 1972. Ixodid ticks (Acarina: Ixodidae) parasitizing wild birds in the Kyasanur Forest disease area of Shimoga District, Mysore State, India. J. Bombay Nat. Hist. Soc., 69: 55–78.

Rajagopalan, P.K. & Sreenivasan, M.A. 1981. Ixodid ticks on cattle and buffaloes in the Kyasanur Forest disease area of Karnataka State. Ind. J. Med. Res., 73: 880–889.

Rajagopalan, P.K., Patil, A.P. & Boshell, J. 1968. Ixodid ticks on their mammalian hosts in the Kyasanur Forest disease area of Mysore State, India, 1961–1964. Ind. J. Med. Res., 56: 510–526.

Warburton, C. 1910. On two collections of Indian ticks. Parasitology, 3: 395–407.

35 – H. dangi Phan Trong, 1977 (Ve bet va con trung ky sinh o Viet Nam. Tap 1. Ve (Ixodoidea), mo ta va phan loai. Ha Noi: Khoa hoc va ky thuat, 489 pp. In Vietnamese)

Type depository: AI (holotype, paratypes) (Kolonin, G.V. personal communication to Guglielmone, A.A.)

Known stages: male, female

Zoogeographic Region: Oriental

Ecoregion: south China-Vietnam subtropical evergreen forests

Hosts: Artiodactyla (several families); Carnivora: Mustelidae; Rodentia: Hystricidae (A)

Human infestation: no

References

Kolonin, G.V. 1995. Review of the ixodid tick fauna (Acari: Ixodidae) of Vietnam. J. Med. Entomol., 32: 276–282.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

36 – H. danieli Černý & Hoogstraal, 1977 (J. Parasitol., 63: 567–574)

Specimens of H. danieli from China are considered identical to H. pospelovashtromae by Kolonin (2009, op. cit. under H. anomaloceraea), but no conclusive evidence is presented. See also “Remarks on some invalid names” at the beginning of this chapter for justification of the status of H. xinjiangensis as a junior synonym of H. danieli, as well as the remarks below.

Type depository: IPCAS (holotype) (Černý, V. & Hoogstraal, H. 1977. Haemaphysalis (Allophysalis) danieli, sp. n., (Ixodoidea: Ixodidae), female and tentatively associated immature stages from high mountains of northern Pakistan and Afghanistan. J. Parasitol., 63: 567–574)

Known stages: male, female, nymph, larva

Zoogeographic Regions: Oriental, Palearctic

Ecoregions: montane grasslands and shrublands in the Oriental and Palearctic Zoogeographic Regions

Hosts: Artiodactyla: Bovidae (A)

Lagomorpha: Ochotonidae; Rodentia: Cricetidae, Muridae, Sciuridae (NL)

Human infestation: no

Remarks: Camicas et al. (1998) state that this species is found only in the Oriental Region, but there are localities included in the original description of H. danieli and in Teng and Jiang (1991), Yu et al. (1997) and Chen et al. (2010) that fall within the Palearctic Region. Camicas et al. (1998) list the male of H. danieli as undescribed. This statement is probably a consequence of regarding H. xinjiangensis as a species separate from H. danieli when in fact the converse is true (see the first paragraph of this chapter). The male of H. danieli was described by Teng (1980), but under the name H. xinjiangensis.

References

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. 1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.

Černý, V. & Hoogstraal, H. 1977. Haemaphysalis (Allophysalis) danieli, sp. n., (Ixodoidea: Ixodidae), female and tentatively associated immature stages from high mountains of northern Pakistan and Afghanistan. J. Parasitol., 63: 567–574.

Chen, Z., Yang, X., Bu, F., Yang, X., Yang, X. & Liu, J. 2010. Ticks (Acari: Ixodoidea: Argasidae, Ixodidae) of China. Exp. Appl. Acarol., 51: 393–404.

Teng, K. 1980. On the Chinese Haemaphysalis subgenus Allophysalis with description of a new species. Acta Entomol. Sin., 23: 86–89. In Chinese.

Teng, K.-F. & Jiang, Z.-J. 1991. Economic insect fauna of China. Fasc. 39, Acari: Ixodidae. Science Press, Beijing, 355 pp. In Chinese.

Yu, X., Ye, R.-E. & Gong, Z.-D. 1997. The tick fauna of Xinjiang. Technological and Medical Publishing House, Urumqi, China, 168 pp. In Chinese.

37 – H. darjeeling Hoogstraal & Dhanda, 1970 (J. Parasitol., 56: 169–174)

Type depositories: BMNH (holotype, paratypes), USNTC, HH, IM (paratypes) (Keirans and Hillyard 2001, op. cit. under H. aciculifer). The specimens used in the description of H. darjeeling were part of the syntype series of H. birmaniae.

Known stages: male, female

Zoogeographic Region: Oriental

Ecoregions: montane grasslands and shrublands

Hosts: Artiodactyla: Bovidae, Cervidae, Suidae; Carnivora: Mustelidae (A)

Human infestation: yes (Tanskul and Inlao 1989, see below)

Remarks: Kolonin (2009) does not include Carnivora as hosts of H. darjeeling, probably because he regards the ticks found on these hosts in Tanskul and Inlao (1989) as closer to H. quadriaculeata than to H. darjeeling, as discussed in Kolonin (2003). We consider the records of this tick on Carnivora in Tanskul et al. (1983) and Tanskul and Inlao (1989) to be provisionally valid. See also H. quadriaculeata.

References

Hoogstraal, H. & Dhanda, V, 1970. Haemaphysalis (H.) darjeeling sp. n., a member of the H. (H.) birmaniae group (Ixodoidea, Ixodidae) parasitizing artiodactyl mammals in Himalayan forests of India, and in Burma and Thailand. J. Parasitol., 56: 169–174.

Kolonin, G.V. 2003. New data on ixodid tick fauna of Vietnam. Entomol. Rev., 83 (Suppl. 2): S190-S192.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Tanskul, P. & Inlao, I. 1989. Keys to the adult ticks of Haemaphysalis Koch, 1844, in Thailand with notes on changes in taxonomy (Acari: Ixodoidea: Ixodidae). J. Med. Entomol., 26: 573–601.

Tanskul, P., Stark, H.E. & Inlao, I. 1983. A checklist of ticks of Thailand (Acari: Metastigmata: Ixodoidea). J. Med. Entomol., 20: 330–341.

38 – H. davisi Hoogstraal, Dhanda & Bhat, 1970 (J. Parasitol., 56: 588–595)

See remarks below.

Type depositories: USNTC (holotype, paratypes), BMNH, VRC, HH, IM (paratypes) (Keirans and Hillyard 2001, op. cit. under H. aciculifer)

Known stages: male, female, nymph

Zoogeographic Region: Oriental

Ecoregion: temperate coniferous forests

Hosts: Artiodactyla: Bovidae, Cervidae; Perissodactyla: Equidae (AN)

Carnivora: Felidae, Mustelidae (A)

Human infestation: no

Remarks: specimens of H. neumanni Dönitz, 1905 (a synonym of Haemaphysalis longicornis) in Dhanda and Rao (1964) are in fact H. davisi (Hoogstraal et al. 1970). Camicas et al. (1998) list Aves and pholeophilic (burrowing) mammals as hosts for larvae and nymphs of this species, but we have found no references to support this statement.

References

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. 1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.

Dhanda, V. & Rao, T.R. 1964. A report on a collection of ixodid ticks made in the North East Frontier Agency, India. Ind. J. Med. Res., 52: 1139–1153.

Hoogstraal, H., Dhanda, V. & Bhat, H.R. 1970. Haemaphysalis (Kaiseriana) davisi sp. n. (Ixodoidea: Ixodidae), a parasite of domestic and wild mammals in northeastern India, Sikkim, and Burma. J. Parasitol., 56: 588–595.

39 – H. demidovae Emel’yanova, 1978 (In Natural conditions and resources of Prikhusbsugul (Mongolian People’s Republic), O.M. Kozhova (editor). Trudy Sov. Mongol. Kompleks. Khubsugul. Eksp. Irkutsk. Gos. Univ. imeni A.A. Zhdanova, Minist. Vyssh. Spets. Obraz. RSFSR (6): 162–171. In Russian).

Type depository: not stated in the description of this species (Keirans, J.E. & Robbins, R.G. 1999. A world checklist of genera, subgenera, and species of ticks (Acari: Ixodida) published from 1973 to 1997. J. Vector Ecol., 24: 115–129).

Known stages: nymph, larva

Zoogeographic Region: Palearctic

Ecoregions: deserts and xeric shrublands

Hosts: Rodentia: Cricetidae, Ochotonidae, Sciuridae (NL)

Human infestation: no

Reference

Emel’yanova, N.D. 1978. Two ixodid species of the subgenus Allophysalis (Haemaphysalis, Ixodidae) from Mongolian People’s Republic. In O.M. Kozhova (editor), Natural conditions and resources of Prikhusbsugul (Mongolian People’s Republic). Trudy Sov. Mongol. Kompleks. Khubsugul. Eksp. Irkutsk. Gos. Univ. Imeni A.A. Zhdanova, Minist. Vyssh. Spets. Obraz. RSFSR (6): 162–171. In Russian, NAMRU-3 translation 1738.

40 – H. doenitzi Warburton & Nuttall, 1909 (Parasitology, 2: 57–76)

See H. phasiana and H. pavlovskyi and remarks below.

Type depository: BMNH (lectotype, paralectotypes) (Keirans and Hillyard 2001, op. cit. under H. aciculifer)

Known stages: male, female, nymph, larva

Zoogeographic Regions: Australasian, Oriental, Palearctic

Ecoregions: tropical and subtropical rain forests

Hosts: usual hosts for larvae, nymphs and adult ticks are Cuculiformes: Cuculidae, Galliformes: Phasianidae. Mammalia and Testudines are considered exceptional hosts for this tick.

Cuculiformes: Cuculidae; Galliformes: Phasianidae; Passeriformes: Sturnidae; Rodentia: Muridae (ANL)

Falconiformes: Accipitridae, Falconidae; Passeriformes: Sylviidae; Strigiformes: Strigidae; Lagomorpha: Leporidae; Testudines: Geoemydidae (AN)

Passeriformes: Alaudidae (AL)

Coraciiformes: Coraciidae, Upupidae; Gruiformes: Rallidae; Passeriformes: Pycnonotidae (A)

Passeriformes: Pittidae, Timaliidae, Turdidae (NL)

Passeriformes: Hirundinidae, Muscicapidae; Piciformes: Capitonidae (N)

Human infestation: yes (Tanskul et al. 1983)

Remarks: although Aves are the preferred hosts of H. doenitzi, Kolonin (2008) believes that Mammalia are equally important as hosts. Kolonin (2009) claims that H. phasiana and H. pavlovskyi are synonyms of H. doenitzi, and perhaps these synonymies, which we consider invalid, are the source of Kolonin’s unexplained statement concerning the host preferences of H. doenitzi. On the other hand, Sames et al. (2008) believe that a complex of species may be represented under the name H. doenitzi. Until this problem is solved, all the above names should be treated as valid. It is not clear whether or not Liu et al. (2002) found specimens of H. doenitzi on Soricidae, and we have therefore excluded this host group from our list above.

References

Hoogstraal, H. & Wassef, H.Y. 1973. The Haemaphysalis ticks (Ixodoidea: Ixodidae) of birds. 3.H. (Ornithophysalis) subgen. n.: definition, species, hosts and distribution in the Oriental, Palearctic, Malagasy, and Ethiopian Faunal Regions. J. Parasitol., 59: 1099–1117.

Kolonin, G.V. 2008. Birds as hosts of ixodid ticks (Acarina: Ixodidae). Entomol. Rev., 88: 1012–1015.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Liu, M.Z., Chen, W,.S. & Gu, W.Z. et al. [sic] 2002. Investigation on rats and insects on their body surface in Shenzhen. Chin. J. Vector Biol. Contr., 13: 13–15. In Chinese.

Sames, W.J., Kim, H.C., Chong, S.T., Lee, I.Y., Apanaskevich, D.A., Robbins, R.G., Bast. J., Moore, R. & Klein, T.A. 2008. Haemaphysalis (Ornithophysalis) phasiana (Acari: Ixodidae) in the Republic of Korea: two province records and habitat descriptions. Syst. Appl. Acarol., 13: 43–50.

Tanskul, P., Stark, H.E. & Inlao, I. 1983. A checklist of ticks of Thailand (Acari: Metastigmata: Ixodoidea). J. Med. Entomol., 20: 330–341.

41 – H. elliptica (Koch, 1844) (Arch. Naturgesch., 10: 217–239)

See remarks below.

Type depository: ZMB (holotype) (Moritz and Fischer 1981, op. cit. under H. cinnabarina) as Rhipistoma ellipticum

Known stages: male, female, nymph, larva

Zoogeographic Region: Afrotropical

Ecoregions: several Afrotropical ecoregions

Hosts: usual hosts for adult ticks are Carnivora: Canidae and Felidae. Aves are considered exceptional hosts for this tick.

Carnivora: Canidae, Felidae (ANL)

Mammalia (several orders) (A)

Macroscelidea: Macroscelididae (N)

Rodentia: Muridae (NL)

Passeriformes: Cisticolidae, Ploceidae (N and/or L)

Human infestation: yes (Apanaskevich et al. 2007)

Remarks: this species has frequently been confused with H. leachi (Apanaskevich et al. 2007). Camicas et al. (1998) listed the male and larva as the only stages of H. elliptica that had been described at the time of their publication; however, the female and nymph were subsequently described by Apanaskevich et al. (2007). The latter authors also indicated that subadults of H. elliptica feed on rodents, without identifying the families, whereas Matthee et al. (2010) collected the immature stages from Muridae. Hasle et al. (2009) do not state whether larvae, nymphs or both immature stages of H. elliptica were found on Passeriformes.

References

Apanaskevich, D.A., Horak, I.G. & Camicas, J.-L. 2007. Redescription of Haemaphysalis (Rhipistoma) elliptica (Koch, 1844), an old taxon of the Haemaphysalis (Rhipistoma) leachi group from East and southern Africa, and of Haemaphysalis (Rhipistoma) leachi (Audouin, 1826) (Ixodida: Ixodidae). Onderstepoort J. Vet. Res., 74: 181–208.

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. 1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.

De Matos, C., Sitoe, C., Neves, L., Bryson, N.R. & Horak, I.G. 2008. Ixodid ticks on dogs belonging to people in rural communities and villages in Maputo Province, Mozambique. Onderstepoort J. Vet. Res., 75: 103–108.

Hasle, G., Horak, I.G., Grieve, G., Leinaas, H.P. & Clarke, F. 2009. Ticks collected from birds in the northern provinces of South Africa, 2004–2006. Onderstepoort J. Vet. Res., 76: 167–175.

Horak, I.G., Heyne, H. & Donkin, E.F. 2010. Parasites of domestic and wild animals in South Africa. XLVIII. Ticks (Acari: Ixodidae) infesting domestic cats and wild felids in southern Africa. Onderstepoort J. Vet. Res., 77, doi:10.4102/ojvr.v77i1.3

Horak, I.G., Welman, S., Hallam, S.L., Lutermann, H. & Mzilikazi, N. 2011. Ticks of four-toed elephant shrews and Southern African hedgehogs. Onderstepoort J. Vet. Res., 78, doi:10.4102/ojvr.v78i1.243

Matthee, S., Horak, I.G., Van der Mescht, L., Ueckermann, E.A. & Radloff. F.G.T. 2010. Ectoparasite diversity on rodents at De Hoop Nature Reserve, Western Cape Province. Afr. Zool., 45: 213–224.

Nyangiwe, N., Goni, S., Hervé-Claude L.P., Ruddat, I. & Horak, I.G. 2012. Ticks on pasture and on two breeds of cattle in the Eastern Cape Province, South Africa. Onderstepoort J. Vet. Res., 78, doi:10.4102/ojvr.v78i1.320

42 – H. elongata Neumann, 1897 (Mém. Soc. Zool. Fr., 10: 324–420)

Type depository: BMNH (syntypes) (Keirans and Hillyard 2001, op. cit. under H. aciculifer)

Known stages: male, female, nymph, larva

Zoogeographic Region: Afrotropical

Ecoregion: Madagascar subhumid forests

Hosts: usual hosts for adult ticks are Afrosoricida: Tenricidae.

Afrosoricida: Tenrecidae (ANL)

Rodentia: Muridae (A)

Carnivora: Eupleridae; Erinaceomorpha: Erinaceidae (N)

Human infestation: yes (Burridge 2011)

Remarks: Uilenberg et al. (1979) regard a record of this tick from Chiroptera as doubtful and we have therefore excluded this record from our list for H. elongata. Keirans and Durden (2001) cite an introduction of this species into the Nearctic Region, but H. elongata has not become established there. Kolonin (2009) and Burridge (2011) ignore Erinaceidae as hosts of this tick, but we consider the record of Keirans (1985) on this host valid.

References

Burridge, M.J. 2011. Non-native and invasive ticks. Threats to human and animal health in the United States. University of Florida Press, Gainesville, 448 pp.

Keirans, J.E. 1985. George Henry Falkiner Nuttall and the Nuttall tick catalogue. U. S. Dept. Agric., Agric. Res. Ser. Misc. Pub. (1438), 1785 pp.

Keirans, J.E. & Durden, L.A. 2001. Invasion: exotic ticks (Acari: Argasidae, Ixodidae) imported into the United States. A review and new records. J. Med. Entomol., 38: 850–861.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Uilenberg, G., Hoogstraal, H. & Klein, J.-M. 1979. Les tiques (Ixodoidea) de Madagascar et leur rôle vecteur. Arch. Inst. Pasteur Madagascar Num. Spéc., 153 pp.

43 – H. erinacei Pavesi, 1884 (Ann. Mus. Civ. Stor. Nat. Genova, 20: 446–486)

The species H. erinacei described in Feldman-Muhsam, B. (1951. A note on east Mediterranean species of the genus Haemaphysalis. Bull. Res. Council Israel, 1: 96–107) is a synonym of H. erinacei Pavesi, 1884, as stated in Hoogstraal, H. (1955. Notes on African Haemaphysalis ticks. I. The Mediterranean-littoral hedgehog parasite. H. erinacei Pavesi, 1884 (Ixodoidea, Ixodidae). J. Parasitol., 41: 221–233)

Many workers believe that this species comprises several subspecies, indicating that more than one taxon is probably included under the name H. erinacei. Filippova, N.A., Panova, I.V. & Musatov, S.A. (1993. Taxonomic structure of the polytypic species Haemaphysalis erinacei (Ixodidae). Parazitologiya, 27: 193–215. In Russian) provide a good description of the subspecies within H. erinacei. See “remarks on some invalid names” for an additional synonym of H. erinacei. See also remarks below.

Type depository: GM (syntypes) (Hoogstraal 1955, op. cit. above)

Known stages: male, female, nymph, larva

Zoogeographic Region: Palearctic

Ecoregions: mediterranean forests, woodlands and scrub

Hosts: Testudines are considered exceptional hosts for this tick.

Carnivora: Canidae; Erinaceomorpha: Erinaceidae (ANL)

Carnivora: Felidae, Hyaenidae, Mustelidae; Lagomorpha: Leporidae, Ochotonidae; Coraciiformes: Meropodidae; Testudines: Testudinidae (A)

Rodentia: Muridae (NL)

Rodentia: Gliridae; Apodiformes: Apodidae; Passeriformes: Motacillidae (N)

Rodentia: Dipodidae (L)

Chiroptera: Vespertilionidae; Rodentia: Calomyscidae, Cricetidae (N and/or L)

Artiodactyla: Bovidae; Carnivora: Ursidae; Rodentia: Sciuridae; Falconiformes: Falconidae; Passeriformes: Emberizidae, Muscicapidae (stages unknown)

Human infestation: yes (Bursali et al. 2012)

Remarks: Serdjukova (1956), Ushakova (1958), Gusev et al. (1961), Bakhaeva (1962), Grebenyuk (1966), and Filippova et al. (1976) call this species H. numidiana. We were unable to determine which immature stages were found by Filippova et al. (1976) on Chiroptera and Rodentia (Calomyscidae and Cricetidae). Tovornik and Černý (1974) doubt whether adult ticks of H. erinacei were found on Coraciiformes (Meropodidae) or Gruiformes (Gruidae), but Tovornik (1990) clearly states that a male was found on Meropodidae. Kolonin (2009) reduces the hosts of H. erinacei to “hedgehogs, small carnivores and rodents” but does not comment on this limited host range.

References

Bakhaeva, A.V. 1962. The fauna of ticks in Turkmenia. Izv. Akad. Nauk Turkmen. Ser. Biol. Nauk. (5): 72–79. In Russian.

Bursali, A., Keskin, A. & Tekin, S. 2012. A review of the ticks (Acari: Ixodida) of Turkey: species diversity, hosts and geographical distribution. Exp. Appl. Acarol., 57: 91–104.

Chen, Z., Yang, X., Bu, F., Yang, X., Yang, X. & Liu, J. 2010. Ticks (Acari: Ixodoidea: Argasidae, Ixodidae) of China. Exp. Appl. Acarol., 51: 393–404.

Filippova, N.A., Neronov, V.M. & Farhang-Azad, A. 1976. Materials on the fauna of ixodids (Acarina, Ixodidae) of small mammals in Iran. Entomol. Obozr., 55: 467–478. In Russian.

Grebenyuk, R.V. 1966. Ixodid ticks (Parasitiformes, Ixodidae) of Kirgizia. Akademii Nauk Kirgizkoy SSR, Instituta Biologii, Frunze, 328 pp. In Russian.

Gusev, V.M., Bednyy, S.N., Guseva, A.A., Labunets, N.F. & Bakeyev, N.N. 1961. The ecological groups of birds on the Caucasus and their role in the life cycle of ticks and fleas. Trudy Nauch.-Issled. Protiv. Inst. Kavk. Zakav. (5): 217–267. In Russian.

Hoogstraal, H. 1955. Notes on African Haemaphysalis ticks. I. The Mediterranean-littoral hedgehog parasite H. erinacei Pavesi, 1884 (Ixodoidea, Ixodidae). J. Parasitol., 41: 221–233.

Hoogstraal, H. 1959. Biological observations on certain Turkish Haemaphysalis ticks (Ixodoidea, Ixodidae). J. Parasitol., 45: 227–232.

Hoogstraal, H., Wassef, H.Y. & Büttiker, W. 1981. Ticks (Acarina) of Saudi Arabia Fam. Argasidae, Ixodidae. Fauna Saudi Arabia, 3: 25–110.

Kolonin, G.V. 1995. Review of the ixodid tick fauna (Acari: Ixodidae) of Vietnam. J. Med. Entomol., 32: 276–282.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Morel, P.-C. 2003. Les tiques d’Afrique et du Bassin méditerranéen (1965–1995). CIRAD- EMVT, 1342 pp.

Serdjukova, G.V. 1956. Ixodid ticks of the fauna of USSR. Opred. Faune SSSR. Zool. Inst. Akad. Nauk SSSR (64), 121 pp. In Russian.

Široký, P., Petrželková, K.J., Kamler, D., Mihalca, A.D. & Modrý, D. 2006. Hyalomma aegyptium as dominant tick in tortoises of the genus Testudo in Balkan countries, with notes on its host preferences. Exp. Appl. Acarol., 40: 279–290.

Theodor, O. & Costa, M. 1967. A survey of the parasites of wild mammals and birds in Israel. Part one. Ectoparasites. The Israel Academy of Science and Humanities, Jerusalem, 119 pp.

Tovornik, D. 1990. The significance of the birds (Aves) as the hosts and disseminators of ixodid ticks (Yugoslavia). Biol. Vestn., 38: 77–108.

Tovornik, D. & Černý, V. 1974. Finding of Haemaphysalis erinacei erinacei Pavesi, 1884 on birds in Yugoslavia. Folia Parasitol., 21: 282.

Ushakova, G.V. 1958. Ixodoidea parasitic on birds of the lower Ili River. Trudy Inst. Zool. Akad. Nauk Kazakh. SSR, 9: 135–145. In Russian.

Ushakova, G.V. & Busalaeva, N.N. 1962. Data on ixodid ticks in the semideserts of the Karaganda Region. Parazit. Sel.-Khoz. Zhivot. Kazakh., 1: 216–224. In Russian.

Wassef, H.I., Büttiker, W. & Gallagher, M.D. 1997. Further records of ticks (Acari: Argasidae and Ixodidae) from the Arab Peninsula. Fauna Saudi Arabia, 16: 63–88.

44 – H. eupleres Hoogstraal, Kohls & Trapido, 1965 (J. Parasitol., 51: 997–1000)

Type depository: AMNH (holotype) (Hoogstraal, H. & Camicas, J.-L. 1977. Haemaphysalis (Rhipistoma) eupleres (Ixodoidea: Ixodidae), a parasite of the Madagascar falanouc (mongoose): new data and male identity. J. Parasitol., 63: 1099–1102). Note: the Madagascar falanouc belongs to the family Eupleridae while mongoose are classified in the family Herpestidae.

Known stages: male, female

Zoogeographic Region: Afrotropical

Ecoregion: Madagascar lowland forests

Hosts: Carnivora: Eupleridae (A)

Human infestation: no

Reference

Uilenberg, G., Hoogstraal, H. & Klein, J.-M. 1979. Les tiques (Ixodoidea) de Madagascar et leur rôle vecteur. Arch. Inst. Pasteur Madagascar Num. Spéc., 153 pp.

45 – H. filippovae Bolotin, 1979 (Zool. Zh., 58: 267–269. In Russian)

Camicas et. al. (1998, op. cit. under H. anomaloceraea) and Kolonin (2009, op. cit. under H. anomaloceraea) treat this species as an abnormal form of H. concinna without further comment. Guglielmone et al. (2009, op. cit. under H. colasbelcouri) consider H. filippovae a valid species and we agree.

Type depository: ZIAC (holotype) (Filippova 2008, op. cit. under H. caucasica)

Known stages: female

Zoogeographic Region: Palearctic

Ecoregions: Ussuri broadleaf and mixed forests

Hosts: unknown

Human infestation: no

References

Bolotin, E.I. 1979. A new Haemaphysalis species (Parasitiformes, Ixodidae) from southern Far East. Zool. Zh., 58: 267–269. In Russian, NAMRU-3 translation 1438.

Filippova, N.A. 1997. Ixodid ticks of subfamily Amblyomminae. Fauna of Russia and neighbouring countries, 4 (5), Nauka, St. Petersburg, 436 pp. In Russian.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

46 – H. flava Neumann, 1897 (Mém. Soc. Zool. Fr., 10: 324–420)

Type depository: MNHN (syntypes) (Neumann, L.G. 1897. Révision de la famille des ixodidés (2^e mémoire). Mém. Soc. Zool. Fr., 10: 324–420)

Known stages: male, female, nymph, larva

Zoogeographic Regions: Oriental, Palearctic

Ecoregions: temperate broadleaf and mixed forests

Hosts: Aves are considered exceptional hosts for adults of H. flava.

Mammalia (several orders); Galliformes: Phasianidae (ANL)

Falconiformes: Accipitridae (A)

Aves (several orders) (NL)

Human infestation: yes (Yamauchi et al. 2010)

Remarks: Camicas et al. (1998) exclude Aves as hosts for the larva and nymph of H. flava, but Miyamoto et al. (2000) subsequently reported the importance of Aves as hosts of the sub-adults of this tick. On the other hand, Kolonin (2009) does not recognize Aves as hosts for adult H. flava, but we accept the record of adults of this species on Galliformes, as reported by Teng and Jiang (1991). See also H. montgomeryi.

References

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. 1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.

Chen, Z., Yang, X., Bu, F., Yang, X., Yang, X. & Liu, J. 2010. Ticks (Acari: Ixodoidea: Argasidae, Ixodidae) of China. Exp. Appl. Acarol., 51: 393–404.

Fujimoto, K., Yamaguti, N. & Takahashi, M. 1986. Ecological studies on ixodid ticks. 1. Ixodid ticks on vegetations and wild animals at the low mountain zone lying south-western part of Saitama Prefecture. Jap. J. Sanit. Zool., 37: 325–331. In Japanese.

Kang, J.-G., Kim, H.-C., Choi, C.-Y., Nam, H.-Y., Chae, H.-Y., Chong, S.-T., Klein, T.A., Ko, S. & Chae, J.-S. 2013. Molecular detection of Anaplasma, Bartonella, and Borrelia species in ticks collected from migratory birds from Hong-do Island, Republic of Korea. Vector- Borne Zoon. Dis., 13: 215–225.

Kobayashi, S., Nakamoto, A., Shimizu, K., Takada, A. & Moriwaki, M. 2012. The first capture with ectoparasitic ticks and the first photograph in life of wild raccon, Procyon lotor (Linnaeus 1758), in Okayama Prefecture. Naturalistae, 16: 83–90. In Japanese.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Kolonin, G.V. & Bolotin, E.I. 1975. New findings of Haemaphysalis flava tick (Parasitioformes, Ixodidae) in the Soviet Primor’e region. Zool. Zh. 54: 616–617. In Russian, NAMRU-3 translation 1274.

Miyamoto, K., Masuzawa, T. & Kudeken, M. 2000. Tick collection from wild birds and detection of Lyme disease spirochetes from a new avian reservoir in Japan. Med. Entomol. Zool., 51: 221–226.

Teng, K.-F. & Jiang, Z.-J. 1991. Economic insect fauna of China. Fasc. 39, Acari: Ixodidae. Science Press, Beijing, 355 pp. In Chinese.

Yamauchi, T. 2001. A bibliographical survey of host-parasite relationships between birds and ticks from Japan. Bull. Hoshizaki Green Found., 5: 271–308. In Japanese.

Yamauchi, T. & Mori, S. 2004. Preliminary study on the tick (Acari: Ixodidae) infestation of wild birds in the eastern part of Shimane Prefecture, Japan. Med. Entomol. Zool., 55: 329–332.

Yamauchi, T., Fukui, Y., Watanabe, M., Nakagawa, H. & Kamimura, K. 2010. Forty cases of human infestations with hard ticks (Acari: Ixodidae) in Toyama Prefecture, Japan. Med. Entomol. Zool., 61: 133–143. In Japanese.

Yamauchi, T., Agetsuma, N., Araki, A. & Fukushima, M. 2012. Ixodid ticks collected from the raccoon dog Nyctereutes procyonoides albus and the common raccoon Procyon lotor in southern Hokkaido, Japan. Int. J. Acarol., 38: 214–216.

Yamauchi, T., Yano, S., Yamamoto, T., Yamamoto, E. & Miyamoto, T. 2013. Ticks (Acari: Ixodidae) from medium-sized to large mammals in Ehime Prefecture, Japan. Exp. Appl. Acarol., 60: 263–270.

47 – H. formosensis Neumann, 1913 (Entomol. Mitt., Suppl. Entomol., 2: 134–137)

Type depositories: BMNH, ZMB (syntypes) (Keirans and Hillyard 2001, op. cit. under H. aciculifer), but its presence in ZMB is not mentionated in Moritz and Fischer (1981, op. cit. under H. cinnabarina)

Known stages: male, female, nymph, larva

Zoogeographic Regions: Oriental, Palearctic

Ecoregions: tropical and subtropical coniferous forests

Hosts: Carnivora: Canidae; Lagomorpha: Leporidae (ANL)

Artiodactyla: Cervidae, Suidae; Galliformes: Phasianidae (AN)

Artiodactyla: Bovidae, Moschidae; Carnivora: Ursidae; Rodentia: Hystricidae, Muridae (A)

Carnivora: Herpestidae; Charadriiformes: Scolopacidae (NL)

Carnivora: Mustelidae; Chiroptera: Vespertilionidae; Scandentia: Tupaiidae (N)

Passeriformes: Corvidae (L)

Human infestation: no

Remarks: Camicas et al. (1998) list the larva and nymph of H. formosensis as undescribed, but Teng and Jiang (1991) had previously provided descriptions of both stages. Kolonin (2009) limits the hosts of H. formosensis to pigs, dogs and muntjacs but provides no explanation for this reduced host range.

References

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. 1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.

Chen, Z., Yang, X., Bu, F., Yang, X., Yang, X. & Liu, J. 2010. Ticks (Acari: Ixodoidea: Argasidae, Ixodidae) of China. Exp. Appl. Acarol., 51: 393–404.

Dhanda, V. & Rao, T.R. 1964. A report on a collection of ixodid ticks made in the North East Frontier Agency, India. Ind. J. Med. Res., 52: 1139–1153.

Ishibashi, O., Niizuma, J., Miura, A., Iizuka, S., Fujita, H., Ogura, G., Sakashita, M., Ganeko, H., Sunagawa, K. & Nakada, T. 2009. Survey of parasitic ixodid ticks on small Asian mongoose on Okinawajima Island, Japan. Jap. J. Zoo Wild. Med., 14: 51–57.

Keirans, J.E. 1985. George Henry Falkiner Nuttall and the Nuttall tick catalogue. U. S. Dept. Agric., Agric. Res. Ser. Misc. Pub. (1438), 1785 pp.

Kitaoka, S. & Suzuki, H. 1974. Reports of medico-zoology investigations in the Nansei Islands. Part 2. Ticks and their seasonal prevalences in southern Amami-oshima. Jap. J. Sanit. Zool., 26: 21–26. In Japanese.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Pang, D., Chen, C. & Xiang, R. 1982. Notes on ixodid ticks from southwestern Yunnan with description of a new species of Haemaphysalis (Acarina: Ixodidae). Zool. Res., 3 (Suppl.): 45–51. In Chinese.

Phan Trong, C. 1977. Ve bet va con trung ky sinh o Viet Nam. Tap 1. Ve (Ixodoidea), mo ta va phan loai. Ha Noi: Khoa hoc va ky thuat, 489 pp. In Vietnamese.

Sugimoto, M. 1937. Notes on the ticks in the Formosa Mountain Reservation for the aborigines. J. Centr. Soc. Vet. Med. Tokyo, 50: 303–340. In Japanese, NAMRU-3 translation 11.

Teng, K.-F. & Jiang, Z.-J. 1991. Economic insect fauna of China. Fasc. 39, Acari: Ixodidae. Science Press, Beijing, 355 pp. In Chinese.

Xu, R. & Luo, G. 1998. Ticks parasitic on Muntiacus reevesi and Lepus sinensis in Wuyi Mountains, Fujian, China. Syst. Appl. Acarol. 3: 197.

Yamauchi, T. & Funakoshi, K. 2000. Ticks (Acari: Ixodoidea) from Chiroptera (Mammalia) of the Kyushu mainland, Japan. J. Acarol. Soc. Jap., 9: 51–54.

Yamauchi, T., Yano, S., Yamamoto, T., Yamamoto, E. & Miyamoto, T. 2013. Ticks (Acari: Ixodidae) from medium-sized to large mammals in Ehime Prefecture, Japan. Exp. Appl. Acarol., 60: 263–270.

48 – H. fossae Hoogstraal, 1953 (Bull. Mus. Comp. Zool., 111: 37–113)

Type depositories: USNTC (holotype, paratypes), BMNH, HH, ISM, FMNH, MCZ, OVI, MNHN (paratypes) (Keirans and Hillyard 2001, op. cit. under H. aciculifer)

Known stages: male, female.

Zoogeographic Region: Afrotropical

Ecoregions: Madagascar dry deciduous forests and succulent woodlands

Hosts: usual hosts for adult ticks are Carnivora: Eupleridae.

Carnivora: Eupleridae, Viverridae (A)

Human infestation: no.

Reference

Uilenberg, G., Hoogstraal, H. & Klein, J.-M. 1979. Les tiques (Ixodoidea) de Madagascar et leur rôle vecteur. Arch. Inst. Pasteur Madagascar Num. Spéc., 153 pp.

49 – H. fujisana Kitaoka, 1970 (Natl. Inst. Anim. Health Q., 10: 73–81)

Type depositories: NIAH (holotype, paratypes), HH, USNTC (paratypes) (Kitaoka, S. 1970. Haemaphysalis (Haemaphysalis) fujisana sp. n. (Ixodoidea, Ixodidae), a cattle parasite in Japan. Natl. Inst. Anim. Health Q., 10: 73–81; Keirans and Clifford 1984, op. cit. under H. bartelsi)

Known stages: male, female, nymph, larva

Zoogeographic Region: Palearctic

Ecoregions: temperate broadleaf and mixed forests

Hosts: Artiodactyla: Bovidae (ANL)

Human infestation: no

Remarks: the male, nymph and larva of H. fujisana are known only from laboratory-reared specimens. Camicas et al. (1998) list pholeophilic (burrowing) mammals as hosts for the larva and nymph, but we have found no records to substantiate this.

References

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. 1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.

Kitaoka, S. 1970. Haemaphysalis (Haemaphysalis) fujisana sp. n. (Ixodoidea, Ixodidae), a cattle parasite in Japan. Natl. Inst. Anim. Health Q., 10: 73–81.

50 – H. garhwalensis Dhanda & Bhat, 1968 (J. Parasitol., 54: 674–678)

Type depositories: VRC (holotype, paratypes), BMNH, IM, HH, USNTC (paratypes) (Keirans and Hillyard 2001, op. cit. under H. aciculifer)

Known stages: male, female, nymph, larva

Zoogeographic Regions: Oriental, Palearctic

Ecoregions: western Himalayan broadleaf forests; Upper Gangetic Plains moist deciduous forests; western Himalayan alpine shrub and meadows

Hosts: usual hosts for adult ticks are Artiodactyla: Bovidae. Rodentia: Cricetidae are considered exceptional hosts for adults of this tick.

Rodentia: Cricetidae (ANL)

Artiodactyla: Bovidae; Perissodactyla: Equidae (A)

Rodentia: Muridae (NL)

Human infestation: no

Remarks: Camicas et al. (1998) list Aves as hosts for larvae and nymphs of this species, but we have been unable to confirm this. Kolonin (2009) ignores Rodentia and Equidae as hosts for adult H. garhwalensis. Although infestation of Muridae by adult H. garhwalensis appears to be an infrequent event, there are sound records for this host-parasite relationship in Dhanda et al. (1977).

References

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. 1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.

Chen, Z., Yang, X., Bu, F., Yang, X., Yang, X. & Liu, J. 2010. Ticks (Acari: Ixodoidea: Argasidae, Ixodidae) of China. Exp. Appl. Acarol., 51: 393–404.

Dhanda, V., Bhat, H.R. & Kulkarni, S.M. 1977. Haemaphysalis (Allophysalis) garhwalensis Dhanda and Bhat (Acarina: Ixodidae), description of immature stages, hosts, distribution and ecology. Ind. J. Parasitol., 1: 9–14.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Teng, K.-F. & Jiang, Z.-J. 1991. Economic insect fauna of China. Fasc. 39, Acari: Ixodidae. Science Press, Beijing, 355 pp. In Chinese.

51 – H. goral Hoogstraal, 1970 (J. Parasitol., 56: 1227–1238)

Type depository: BMNH (holotype) (Keirans and Hillyard 2001, op. cit. under H. aciculifer). This tick was originally classified as H. neumanni Dönitz, 1905 (Keirans, J.E. 1985. George Henry Falkiner Nuttall and the Nuttall tick catalogue. U. S. Dept. Agric., Agric. Res. Ser. Misc. Publ. (1438), 1785 pp), which is a synonym of H. longicornis.

Known stages: female

Zoogeographic Region: Oriental

Ecoregion: Jian Nan subtropical evergreen forests

Hosts: Artiodactyla: Bovidae (A)

Human infestation: no

Remarks: Mihalca et al. (2011) regard H. goral as an endangered species.

References

Chen, Z., Yang, X., Bu, F., Yang, X., Yang, X. & Liu, J. 2010. Ticks (Acari: Ixodoidea: Argasidae, Ixodidae) of China. Exp. Appl. Acarol., 51: 393–404.

Hoogstraal, H. 1970. Haemaphysalis (H.) birmaniae Supino and H. (H.) goral sp. n. (Ixodoidea: Ixodidae), Asian parasites of artiodactyl mammals. J. Parasitol., 56: 1227–1238.

Mihalca, A.D., Gherman, C.M. & Cozma, V. 2011. Coendangered hard-ticks: threatened or threatening? Parasit. Vectors, 4 (71), 7 pp.

52 – H. grochovskajae Kolonin, 1992 (In Sokolov, V.E. (editor), Zoological Researches in Vietnam. Nauka, Moscow, pp. 242–277. In Russian)

Type depositories: ZIAC (holotype), KC (paratype) (Filippova 2008, op. cit. under H. caucasica)

Known stages: female

Zoogeographic Region: Oriental

Ecoregion: northern Indochina subtropical forests

Hosts: Artiodactyla: Bovidae (A)

Human infestation: no

Reference

Kolonin, G.V. 1995. Review of the ixodid tick fauna (Acari: Ixodidae) of Vietnam. J. Med. Entomol., 32: 276–282.

53 – H. heinrichi Schulze, 1939 (Z. Parasitenkd., 10: 722–728)

See remarks below.

Type depository: USNTC (lectotype, paralectotypes) (Keirans and Clifford 1984, op. cit. under H. bartelsi)

Known stages: male, female, nymph

Zoogeographic Region: Oriental

Ecoregions: tropical and subtropical moist broadleaf forests

Hosts: Carnivora: Canidae, Mustelidae (A, N and/or L)

Rodentia: Muridae (AN)

Artiodactyla: Bovidae; Carnivora: Viverridae; Rodentia: Sciuridae (A)

Carnivora: Herpestidae; Soricomorpha: Soricidae (N and/or L)

Human infestation: yes (Hoogstraal and Kohls 1968)

Remarks: Hoogstraal and Kim (1985) state that H. heinrichi is a polytypic species, and therefore it is possible that two or more species are represented under the name H. heinrichi. The host-parasite relationships above will probably change if this statement is confirmed. Tanskul et al. (1983) and Hoogstraal and Kim (1985) use the term “immatures” without stating whether larvae, nymphs or both stages were found on hosts. Tanskul et al. (1983) present records of H. heinrichi adults on Bovidae that we consider provisionally valid. However, this type of host is not mentioned in Kolonin (2009).

References

Hoogstraal, H. & Kim, K.C. 1985. Tick and mammal coevolution, with emphasis on Haemaphysalis. In K.C. Kim (editor), Coevolution of parasitic arthropods and mammals. John Wiley & Sons, New York, pp. 505–568.

Hoogstraal, H. & Kohls, G.M. 1968. Studies on Southeast Asian Haemaphysalis ticks (Ixodoidea, Ixodidae). Redescription of type material of H. (Rhipistoma) heinrichi Schulze, a parasite of the ferret-badger in Burma and Vietnam. J. Parasitol., 54: 1057–1062.

Kolonin, G.V. 2003. New data on ixodid tick fauna of Vietnam. Entomol. Rev., 83 (Suppl. 2): S190-S192.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Phan Trong, C. 1977. Ve bet va con trung ky sinh o Viet Nam. Tap 1. Ve (Ixodoidea), mo ta va phan loai. Ha Noi: Khoa hoc va ky thuat, 489 pp. In Vietnamese.

Tanskul, P., Stark, H.E. & Inlao, I. 1983. A checklist of ticks of Thailand (Acari: Metastigmata: Ixodoidea). J. Med. Entomol., 20: 330–341.

54 – H. hirsuta Hoogstraal, Trapido & Kohls, 1966 (J. Parasitol., 52: 169–191)

Type depository: MCZ (holotype, paratype) (Hoogstraal, H., Trapido, H. & Kohls, G.M. 1966. Studies on Southeast Asian Haemaphysalis ticks (Ixodoidea, Ixodidae). Speciation in the H. (Kaiseriana) obesa group: H. semermis Neumann, H. obesa Larrousse, H. roubaudi Toumanoff, H. montgomeryi Nuttall, and H. hirsuta sp. n. J. Parasitol., 52: 169–191), originally classified as Haemaphysalis papuana.

Known stages: male, female

Zoogeographic Region: Oriental

Ecoregions: Sumatran and Java rain forests

Hosts: Artiodactyla: Bovidae, Suidae; Carnivora: Canidae, Mustelidae; Rodentia: Hystricidae (A)

Artiodactyla: Cervidae; Carnivora: Viverridae (N and/or L)

Human infestation: yes (Hoogstraal and Kim 1985)

Remarks: Hoogstraal and Kim (1985) use the term “immatures” without specifying whether larvae, nymphs or both stages of H. hirsuta were found on hosts. Neither of these stages has formally been described, but we provisionally accept their diagnosis by these authors.

References

Hoogstraal, H. & Kim, K.C. 1985. Tick and mammal coevolution, with emphasis on Haemaphysalis. In K.C. Kim (editor), Coevolution of parasitic arthropods and mammals. John Wiley & Sons, New York, pp. 505–568.

Hoogstraal, H., Trapido, H. & Kohls, G.M. 1966. Studies on Southeast Asian Haemaphysalis ticks (Ixodoidea, Ixodidae). Speciation in the H. (Kaiseriana) group: H. semermis Neumann, H. obesa Larrousse, H. roubaudi Toumanoff, H. montgomeryi Nuttall, and H. hirsuta sp. n. J. Parasitol., 52: 169–191.

Phan Trong, C. 1977. Ve bet va con trung ky sinh o Viet Nam. Tap 1. Ve (Ixodoidea), mo ta va phan loai. Ha Noi: Khoa hoc va ky thuat, 489 pp. In Vietnamese.

55 – H. hispanica Gil Collado, J. 1938 (Brotéria Cienc. Nat., 7: 99–109)

Type depository: CGC-FF (holotype, paratypes) (Hoogstraal, H. & Morel, P.-C. 1970. Haemaphysalis (Rhipistoma) hispanica Gil Collado, a parasite of the European rabbit, redescription of adults, and description of immature stages (Ixodoidea: Ixodidae). J. Parasitol., 56: 813–822) as Haemaphysalis campanulata hispanica. These authors state that the type specimens were in the possession of Professor Gil Collado, but one of us (AEP) located the CGC collection within the FF collection.

Known stages: male, female, nymph, larva

Zoogeographic Region: Palearctic

Ecoregions: Iberian sclerophyllous and semi-deciduous forests

Hosts: usual hosts for adult ticks are Lagomorpha: Leporidae. Aves are considered exceptional hosts for this tick.

Lagomorpha: Leporidae (ANL)

Strigiformes: Strigidae (A)

Falconiformes: Accipitridae (L)

Human infestation: no

Remarks: González Montana et al. (1998) reported heavy infestations of dogs with H. hispanica, but we consider this doubtful. Santos Dias (1994) states that Gil Collado found this species on several orders of mammals, but we have been unable to confirm this. These vertebrates are not included in our host list for H. hispanica. Kolonin (2009) ignores the record of this tick on Aves, but we accept the exceptional record on birds in Estrada-Peña et al. (1985).

References

Estrada-Peña, A., Lucientes Curdi, J., Sánchez Acedo, C., Gutiérrez Galindo, J., Ocabo Meléndez, B., Galmes Femenias, M. & Castillo Hernández, J. 1985. Parasitismo accidental de Haemaphysalis (Rhipistoma) hispanica Gill Collado (Acari: Ixodoidea) sobre Aquila chrysaetos. Rev. Ibér. Parasitol., 45: 379–380.

González Montana, J.R., Rejas López, J., Alonso Diez, A.J., Alonso Alonso, M.P. & Prieto Montana, F. 1998. Utilización de citioato para controlar infestaciones por garrapatas en perros. Med. Vet. (Barcelona), 15: 143–146.

Hoogstraal, H. & Morel, P.-C. 1970. Haemaphysalis (Rhipistoma) hispanica Gil Collado, a parasite of the European rabbit, redescription of adults, and description of immature stages (Ixodoidea: Ixodidae). J. Parasitol., 56: 813–822.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Santos Dias, J.A.T. 1994. As carraças (Acarina: Ixodoidea) da península ibérica. Algumas considerações sobre a sua biogeografía e relacionamento com a ixodofauna afropaleártica e afrotropical. Estud. Ens. Doc. (158), 163 pp.

56 – H. hoodi Warburton & Nuttall, 1909 (Parasitology, 2: 57–76)

Type depository: BMNH (lectotype, paralectotypes) (Keirans and Hillyard 2001, op. cit. under H. aciculifer)

Known stages: male, female, nymph, larva

Zoogeographic Region: Afrotropical

Ecoregions: tropical and subtropical grasslands, savannas and shrublands

Hosts: Mammalia are considered exceptional hosts for this tick.

Aves (several orders) (ANL)

Primates: Indriidae (AN)

Artiodactyla: Bovidae; Rodentia: Sciuridae (A, N and/or L)

Carnivora: Canidae; Lagomorpha: Leporidae; Rodentia: Hystricidae (A)

Carnivora: Viverridae (N and/or L)

Human infestation: no

Remarks: Keirans and Durden (2001) record an introduction of H. hoodi into the Nearctic Region, but there is no evidence that it has become established there. Theiler (1962) uses the term “immatures” without specifying whether larvae, nymphs or both stages of H. hoodi were found on hosts. Infestation of Mammalia by this tick is not a common event, but there are some examples of this host-parasite relationship in the references below. Kolonin (2009) ignores mammals as hosts for H. hoodi but provides no reason doing so.

References

Aeschlimann, A. 1967. Biologie et écologie des tiques (Ixodoidea) de Côte d’Ivoire. Acta Trop., 24: 281–405.

Cornet, J.-P. 1995. Contribution à l’étude des tiques (Acarina: Ixodina) de la République Centrafricaine 4. Inventaire et répartition. Acarologia, 36: 203–212.

Elbl, A. & Anastos, G. 1966. Ixodid ticks (Acarina, Ixodidae) of Central Africa. Vol. IV. Genera Aponomma Neumann, 1899, Boophilus Curtice, 1891, Dermacentor Koch, 1844, Haemaphysalis Koch, 1844, Hyalomma Koch, 1844 and Rhipicentor Nuttall and Warburton, 1908. Ann. Mus. R. Centr. Afr., Ser. 8º Sci. Zool. (148), 412 pp.

Horak, I.G., Spickett, A.M., Braack, L.E.O., Penzhorn, B.L., Bagnall, R.J. & Uys, A.C. 1995. Parasites of domestic and wild animals in South Africa. XXXIII. Ixodid ticks on scrub hares in the north-eastern regions of Northern Transvaal and of KwaZulu-Natal. Onderstepoort J. Vet. Res., 62: 123–131.

Keirans, J.E. 1985. George Henry Falkiner Nuttall and the Nuttall tick catalogue. U. S. Dept. Agric., Agric. Res. Ser. Misc. Pub. (1438), 1785 pp.

Keirans, J.E. & Durden, L.A. 2001. Invasion: exotic ticks (Acari: Argasidae, Ixodidae) imported into the United States. A review and new records. J. Med. Entomol., 38: 850–861.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Matthysse, J.G. & Colbo, M.H. 1987. The ixodid ticks of Uganda. Entomological Society of America, College Park, Maryland, 426 pp.

Santos Dias, J.A.T. 1958. Notes on various ticks (Acarina-Ixodoidea) in collection at some entomological Institutes in Paris and London. An. Inst. Med. Trop., 15: 459–563.

Theiler, G. 1962. The Ixodoidea parasites of vertebrates in Africa south of the Sahara (Ethiopian Region). Report to the Director of Veterinary Services, Onderstepoort, South Africa, Project S.9958, 260 pp.

57 – H. hoogstraali Kohls, 1950 (Natl. Inst. Health Bull. (192), 28 pp)

Type depositories: CNHM (holotype, paratypes), USNTC (paratypes) (Kohls, G.M. 1950. Ticks (Ixodoidea) of the Philippines. Natl. Inst. Health Bull. (192), 28 pp.)

Known stages: male, female

Zoogeographic Region: Oriental

Ecoregion: Palawan rain forests

Hosts: Carnivora: Mephitidae (A)

Human infestation: no

Reference

Kohls, G.M. 1950. Ticks (Ixodoidea) of the Philippines. Natl. Inst. Health Bull. (192), 28 pp.

58 – H. houyi Nuttall & Warburton, 1915 (Ticks. A monograph of the Ixodoidea. Part III. The genus Haemaphysalis. Cambridge University Press, London, pp. 349–550)

Type depositories: BMNH (lectotype, paralectotypes), ZMB (paralectotypes) (Keirans and Hillyard 2001, op. cit. under H. aciculifer), originally named Haemaphysalis calcarata houyi.

Known stages: male, female, nymph, larva

Zoogeographic Region: Afrotropical

Ecoregions: tropical and subtropical grasslands, savannas and shrublands

Hosts: usual hosts for larvae, nymphs and adults are Rodentia: Sciuridae.

Rodentia: Sciuridae (ANL)

Rodentia: Muridae; Soricomorpha: Soricidae (AN)

Erinaceomorpha: Erinaceidae; Carnivora: Mustelidae, Viverridae (A)

Hyracoidea: Procaviidae (stage unknown)

Human infestation: no

Remarks: Kolonin (2009) excludes records of H. houyi on hosts other than Sciuridae, but we accept several of the records from other hosts in the references below.

References

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. 1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.

Cornet, J.-P., Zeller, H., Bâ, K., Camicas, J.-L., González, J.-P. & Wilson, M.L. 1995. Contribution a l’étude des tiques (Acarina: Ixodina) vectrices du virus de la fièvre hémorragique Crimée-Congo (CCHF) au Sénégal. I. Analyse du parasitisme chez les petits rongeurs. Acarologia, 36: 287–292.

Hoogstraal, H. 1955. Notes on African Haemaphysalis ticks. II. The ground-squirrel parasites, H. calcarata Neumann, 1902, and H. houyi Nuttall and Warburton, 1915 (Ixodoidea, Ixodidae). J. Parasitol., 41: 361–373.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Matthysse, J.G. & Colbo, M.H. 1987. The ixodid ticks of Uganda. Entomological Society of America, College Park, Maryland, 426 pp.

Morel, P.-C. 1978. Tiques d’animaux sauvages en Haute-Volta. Rev. Élev. Méd. Vét. Pays Trop., 31: 69–78.

Morel, P.-C. 2003. Les tiques d’Afrique et du Bassin méditerranéen (1965–1995). CIRAD- EMVT, 1342 pp.

Theiler, G. 1962. The Ixodoidea parasites of vertebrates in Africa south of the Sahara (Ethiopian Region). Report to the Director of Veterinary Services, Onderstepoort, South Africa, Project S.9958, 260 pp.

59 – H. howletti Warburton, 1913 (Parasitology, 6: 121–130)

Type depository: BMNH (lectotype, paralectotype) (Keirans and Hillyard, 2001, op. cit. under H. aciculifer)

Known stages: male, female, nymph, larva

Zoogeographic Region: Oriental

Ecoregion: southern Vietnam lowland dry forests

Hosts: Rodentia: Muridae (ANL)

Lagomorpha: Leporidae; Persissodactyla: Equidae (A)

Cuculiformes: Cuculidae (N)

Rodentia: Sciuridae (L)

Carnivora: Felidae, Herpestidae; Galliformes: Phasianidae (stages unknown)

Human infestation: no

Remarks: infestation of Rodentia by a male of H. howletti, as reported in Dhanda (1964), is ignored by Kolonin (2009). Rao et al. (1973) list “mongoose” as a host for H. howletti without further discussion, which is probably why Mitchell (1979) also lists Herpestidae as hosts of H. howletti. As a consequence, we have tentatively included this host in our list.

References

Dhanda, V. 1964. Description of immature stages of Haemaphysalis howletti (Ixodoidea: Ixodidae), and redescription of adults. J. Parasitol., 50: 459–465.

Kolonin, G.V. 1995. Review of the ixodid tick fauna (Acari: Ixodidae) of Vietnam. J. Med. Entomol., 32: 276–282.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Mitchell, R.M. 1979. A list of ectoparasites from Nepalese mammals, collected during the Nepal ectoparasite program. J. Med. Entomol., 16: 227–233.

Mitchell, R.M. & Dick, J.A. 1978. Ectoparasites from Nepal birds. J. Bombay Nat. Hist. Soc., 74: 264–274.

Rao, T.R., Dhanda, V., Bhat, H.R. & Kulkarni, S.M. 1973. A survey of haematophagous arthropods in western Himalayas, Sikkim and Hill Districts of West Bengal. A general account. Ind. J. Med. Res., 61: 1421–1461.

60 – H. humerosa Warburton & Nuttall, 1909 (Parasitology, 2: 57–76)

Type depository: BMNH (lectotype, paralectotypes) (Keirans and Hillyard 2001, op. cit. under H. aciculifer)

Known stages: male, female, nymph, larva

Zoogeographic Region: Australasian

Ecoregions: tropical and subtropical grasslands, savannas and shrublands

Hosts: usual hosts for larvae, nymphs and adult ticks are Peramelemorphia: Peramelidae.

Peramelemorphia: Peramelidae; Rodentia: Muridae (ANL)

Monotremata: Tachyglossidae (A)

Cuculiformes: Cuculidae (NL)

Dasyuromorphia: Dasyuridae (L)

Artiodactyla: Bovidae; Perissodactyla: Equidae; Diprotodontia: Phalangeridae; Coraciiformes: Coraciidae (stages unknown)

Human infestation: yes (Stewart and De Vos 1984)

Remarks: Kolonin (2009) ignores Muridae as hosts for H. humerosa, but there are bona fide records of this host-parasite relationship in Roberts (1963, 1970), and more recently in Weaver and Smales (2012).

References

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Roberts, F.H.S. 1963. A systematic study of the Australian species of the genus Haemaphysalis Koch (Acarina: Ixodidae). Aust. J. Zool., 11: 35–80.

Roberts, F.H.S. 1970. Australian ticks. CSIRO, Melbourne, 267 pp.

Stewart, N.P. & De Vos, A.J. 1984. Ticks and the diseases they carry. Queensland Agric. J., 110: 295–299.

Weaver, H.J. & Smales, L.R. 2012. Parasite assemblage of Australian species of Pseudomys (Rodentia: Muridae: Murinae). J. Parasitol., 98: 30–35.

61 – H. hylobatis Schulze, 1933 (Arch. Hydrobiol. Suppl. 12, 4: 490–502)

Type depositories: ZMB (holotype), USNTC (paratype) (Anastos, G. 1950. The scutate ticks, or Ixodidae, of Indonesia. Entomol. Am., 30: 1–144; Keirans and Clifford 1984, op. cit. under H. bartelsi). Moritz and Fischer (1981, op. cit. under H. cinnabarina) do not confirm the presence of the holotype of H. hylobatis in ZMB.

Known stages: male, female

Zoogeographic Region: Oriental

Ecoregion: Sumatran lowland rain forests

Hosts: Aves are considered exceptional hosts for adults of this tick.

Mammalia (several orders); Cuculiformes: Cuculidae (A)

Rodentia: Muridae (N)

Human infestation: yes (Hoogstraal and Kim 1985)

Remarks: Audy et al. (1960) state that the determination of a nymph of H. hylobatis from Tragulidae is tentative, and this family has therefore been excluded from our host list for this species. However, the record of a nymph from Muridae in Audy et al. (1960) is certain because the nymph molted into a female of H. hylobatis.

References

Anastos, G. 1950. The scutate ticks, or Ixodidae, of Indonesia. Entomol. Am., 30: 1–144.

Audy, J.R., Nadchatram, M. & Lim, B.-L. 1960. Malaysian parasites. XLIX. Host distribution of Malayan ticks (Ixodoidea). Stud. Inst. Med. Res. Malaya, 29: 225–246.

Hoogstraal, H. & Kim, K.C. 1985. Tick and mammal coevolution, with emphasis on Haemaphysalis. In K.C. Kim (editor), Coevolution of parasitic arthropods and mammals. John Wiley & Sons, New York, pp. 505–568.

Hoogstraal, H., Lim, B.L., Nadchatram, M. & Anastos, G. 1972. The Gunong Benom Expedition 1967. 8. Ticks (Ixodidae) of Gunong Benom and their altitudinal distribution, hosts and medical relationships. Bull. Br. Mus. (Nat. Hist.) Zool., 23: 167–186.

62 – H. hyracophila Hoogstraal, Walker & Neitz, 1971 (J. Parasitol., 57: 417–425)

Type depositories: USNTC (holotype, paratypes), BMNH, OVI, SAIMR, HH (paratypes) (Keirans and Hillyard 2001, op. cit. under H. aciculifer)

Known stages: male, female, nymph, larva

Zoogeographic Region: Afrotropical

Ecoregions: rocky outcrops and mountains in tropical and subtropical grasslands, savannas and shrublands

Hosts: usual hosts for larvae, nymphs and adults are Hyracoidea: Procaviidae, while Macroscelidea are considered exceptional hosts for this tick.

Hyracoidea: Procaviidae (ANL)

Macroscelidea: Macroscelididae (NL)

Human infestation: no

Remarks: the nymphs and larvae of H. hyracophila collected from Procaviidae by Horak and Fourie (1986) were tentatively thought to belong to this species, a host-parasite relationship that was later confirmed by Fourie et al. (2005). Kolonin (2009) ignores the bona fide records of larvae and nymphs of H. hyracophila on Macroscelidea reported by Fourie et al. (2005), probably because the latter authors regarded the few ticks found on these hosts as “stragglers.”

References

Fourie, L.J., Horak, I.G. & Woodall, P.F. 2005. Elephant shrews as hosts of immature ixodid ticks. Onderstepoort J. Vet. Res., 72: 293–301.

Horak, I.G. & Fourie, L.J. 1986. Parasites of domestic and wild animals in South Africa. XIX. Ixodid ticks and fleas on rock dassies (Procavia capensis) in the Mountain Zebra National Park. Onderstepoort J. Vet. Res., 53: 123–126.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

63 – H. hystricis Supino, 1897 (Atti Soc. Veneto-Trentina Sci. Nat. Residente Padova, Ser. 2, 3: 230–238)

Type depositories: BMNH (lectotype, paralectotype), GM (paralectotype) (Keirans & Hillyard, 2001, op. cit. under H. aciculifer)

Known stages: male, female, nymph, larva

Zoogeoegraphic Regions: Australasian, Oriental, Palearctic

Ecoregions: tropical and subtropical forests

Hosts: Aves are considered exceptional hosts for this tick.

Mammalia (several orders) (ANL)

Cuculiformes: Cuculidae (L)

Passeriformes (several families) (stages unknown)

Human infestation: yes (Durden et al. 2008)

Remarks: with the exception of the type data, records of this species published prior to Hoogstraal et al. (1965) have been ignored because of diagnostic uncertainties. Camicas et al. (1998) state that this species is found in the Oriental and Palearctic Zoogeographic Regions, but Durden et al. (2008) confirmed its presence in the Australasian Region as well. Camicas et al. (1998) include Testudines as hosts for this species because there are records from Geoemyda spinosa (= Heosemys spinosa) prior to 1965, but we regard these as doubtful and they have not been included in our host list for H. hystricis. These questionable records are repeated in Barnard and Durden (2000). Aves are ignored as hosts of H. hystricis, but without elaboration, in Kolonin (2009). We treat records of H. hystricis on Aves in Jiang (1983) and Yamauchi (2001) as valid.

References

Barnard, S.M. & Durden, L.A. 2000. A Veterinary guide to the parasites of reptiles. Volume 2: Arthropods (Excluding Mites). Malabar, Florida: Krieger Publishing Co. 288 pp.

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. 1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.

Chen, Z., Yang, X., Bu, F., Yang, X., Yang, X. & Liu, J. 2010. Ticks (Acari: Ixodoidea: Argasidae, Ixodidae) of China. Exp. Appl. Acarol., 51: 393–404.

Dilrukshi, P.R.M.P. 2006. Taxonomic status of ticks in Sri Lanka. In C.N.B. Bambaradeniya (editor), The fauna of Sri Lanka: status of taxonomy, research and conservation. The World Conservation Union, Colombo, Sri Lanka, pp. 65–69.

Durden, L.A., Merker, S. & Beati, L. 2008. The tick fauna of Sulawesi, Indonesia (Acari: Ixodoidea: Argasidae and Ixodidae). Exp. App. Acarol., 45: 85–110.

Grassman, L.I., Sarataphan, N., Tewes, M.E., Silvy, N.J. & Nakanakrat, T. 2004. Ticks (Acari: Ixodidae) parasitizing wild carnivores in Phu Khieo wildlife sanctuary. J. Parasitol., 90: 657–659.

Hoogstraal, H., Trapido, H. & Kohls, G.M. 1965. Studies on Southeast Asian Haemaphysalis ticks (Ixodoidea, Ixodidae). The identity, distribution, and hosts of H. (Kaiseriana) hystricis Supino. J. Parasitol., 51: 467–480.

Hoogstraal, H., Santana, F.J. & Van Peenen, P.F.D. 1968. Ticks (Ixodoidea) of Mt. Sontra, Danang, Republic of Vietnam. Ann. Entomol. Soc. Am., 61: 722–729.

Jiang, Z.-J. 1983. Biology of Haemaphysalis hystricis Supino. Acta Entomol. Sin., 11: 413–418. In Chinese.

Kitaoka, S. & Suzuki, H. 1974. Reports of medico-zoology investigations in the Nansei Islands. Part 2. Ticks and their seasonal prevalences in southern Amami-oshima. Jap. J. Sanit. Zool., 26: 21–26. In Japanese.

Senadhira, M.A.P. 1969. The parasites of Ceylon. V. Arthropoda, a host check list. Ceylon Vet. J., 17: 3–25.

Shimada, Y., Inokuma, H., Beppu, T., Okuda, M. & Onishi, T. 2003. Survey of ixodid tick species on domestic cats in Japan. Vet. Parasitol., 111: 231–239.

Supino, F. 1897. Nuovi Ixodes della Birmaina (nota preventiva). Atti Soc. Veneto-Trentina Sci. Nat. Residente Padova, Ser. 2, 3: 230–238.

Tanskul, P. & Inlao, I. 1989. Keys to the adult ticks of Haemaphysalis Koch, 1844, in Thailand with notes on changes in taxonomy (Acari: Ixodoidea: Ixodidae). J. Med. Entomol., 26: 573–601.

Yamaguti, N., Tipton, V.J., Keegan, H.L. & Toshioka, S. 1971. Ticks of Japan, Korea, and the Ryukyu Islands. Brigham Young Univ. Sci. Bull. Biol. Ser., 15 (1), 226 pp.

Yamauchi, T. 2001. A bibliographical survey of host-parasite relationships between birds and ticks from Japan. Bull. Hoshizaki Green Found., 5: 271–308. In Japanese.

64 – H. ias Nakamura & Yajima, 1937 (Rep. Gov. Exp. Stn. Anim. Hyg. Tokyo (17): 133–184. In Japanese)

See remarks below

Type depository: CK (holotype) (Yamaguti, N., Tipton, V.J., Keegan, H.L. & Toshioka, S. (1971. Ticks of Japan, Korea, and the Ryukyu Islands. Brigham Young Univ. Sci. Bull. Biol. Ser., 15 (1), 226 pp.)

Known stages: male, female, nymph, larva

Zoogeographic Region: Palearctic

Ecoregions: temperate broadleaf and mixed forests

Hosts: Artiodactyla: Cervidae (AN)

Artiodactyla: Bovidae (A)

Aves (unknown orders) (NL)

Carnivora: Canidae (N)

Human infestation: no

Remarks: Yamaguti et al. (1971) referred to this species as H. cornigera group, H. cornigera, or H. ias. We regard all such records in Yamaguti et al. (1971) as H. ias. Mori et al. (1995) refer to this tick as H. cornigera ias, while Shimada et al. (2003) refer to it as H. ias. Takada (1990) and Fujita and Takada (2007) excluded the name ias but retained cornigera as a Japanese tick. Further studies or better translations of the Japanese literature are needed in order to assess the validity of these taxa. Heath (2013) lists humans as hosts of H. ias as a result of a transcription error (Heath, A.C.G., personal communication to Guglielmone, A.A.). See also H. cornigera.

References

Fujita, H. & Takada, N. 2007. Identification of immatures ticks in Japan. In Organization Committee of SADI (editor), Acari and Emerging/Reemerging Diseases. Zenkoku Nosom Kyoiku Kyokai Publishing Co, Tokyo, pp. 53–68. In Japanese.

Heath, A.C.G. 2013. Implications for New Zealand of potentially invasive ticks sympatric with Haemaphysalis longicornis Neumann, 1901 (Acari: Ixodidae). Syst. Appl. Acarol., 18: 1–26.

Mori, K., Tsunoda, T. & Fujimagari, M. 1995. Ixodid ticks on Sika deer Cervus nippon Temminck in Chiba Prefecture. Jap. J. Sanit. Zool., 46: 313–316.

Shimada, Y., Beppu, T., Inokuma, H., Okuda, M. & Onishi, T. 2003. Ixodid tick species recovered from domestic dogs in Japan. Med. Vet. Entomol., 17: 38–45.

Takada, N. 1990. A pictorial review of medical acarology in Japan. Kinpodo Press, Kyoto, Japan, 222 pp. In Japanese, translation of the legends of figures by Kore, V.

Yamaguti, N., Tipton, V.J., Keegan, H.L. & Toshioka, S. 1971. Ticks of Japan, Korea, and the Ryukyu Islands. Brigham Young Univ. Sci. Bull. Biol. Ser., 15 (1), 226 pp.

65 – H. indica Warburton, 1910 (Parasitology, 3: 395–407)

See remarks below.

Type depositories: BMNH, IM (syntypes) (Keirans and Hillyard 2001, op. cit. under H. aciculifer) as H. leachi indica

Known stages: male, female, nymph, larva

Zoogeographic Regions: Afrotropical, Oriental, Palearctic

Ecoregions: tropical and subtropical dry broadleaf forests

Hosts: usual hosts for adult ticks are Carnivora: Herpestidae. Aves are considered exceptional hosts for this tick.

Carnivora: Canidae, Felidae, Herpestidae (ANL)

Carnivora: Viverridae; Lagomorpha: Leporidae; Galliformes: Phasianidae (AN)

Carnivora: Mustelidae (AL)

Artiodactyla: Cervidae; Carnivora: Hyaenidae (A)

Soricomorpha: Soricidae (N)

Passeriformes: Muscicapidae, Timaliidae (NL)

Passeriformes: Laniidae (L)

Artiodactyla: Suidae; Erinaceomorpha: Erinaceidae; Rodentia: Muridae (N and/or L)

Cuculiformes: Cuculidae; Passeriformes: Turdidae (stage unknown)

Human infestation: no

Remarks: records published prior to Hoogstraal (1970) have not been included because of diagnostic uncertainties. Camicas et al. (1998) state that this species is found only in the Oriental Region, but there are also records from the Afrotropical and Palearctic Regions. However, the distribution of H. indica should be considered cautiously because Hoogstraal and Kim (1985) state that more than one taxon may be included under this name. These authors assert that H. indica is found in the Oriental and Palearctic Zoogeographic Regions, but mention its presence also in Oman, which we regard as part of the Afrotropical Region. Kaul et al. (1978), Hoogstraal (1980) and Geevarghese and Dhanda (1995) do not specify whether they found larvae, nymphs or both immature stages of I. indica on hosts. Dilrukshi (2006) provides records from Bovidae, Sciuridae and Aves that he classifies as H. leachi indica, but these have not been included in our host list because we believe that they need confirmation. Kolonin (2009) excludes Aves as hosts of H. indica, but we consider the records for this group of hosts in Kaul et al. (1978) to be valid.

References

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. 1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.

Dilrukshi, P.R.M.P. 2006. Taxonomic status of ticks in Sri Lanka. In C.N.B. Bambaradeniya (editor), The fauna of Sri Lanka: status of taxonomy, research and conservation. The World Conservation Union, Colombo, Sri Lanka, pp. 65–69.

Geevarghese, G. & Dhanda, V. 1995. Ixodid ticks of Maharashtra State, India. Acarologia, 36: 309–313.

Hoogstraal, H. 1970. Identity, distribution, and hosts of Haemaphysalis (Rhipistoma) indica Warburton (resurrected) (Ixodoidea: Ixodidae), a carnivore parasite of the Indian subregion. J. Parasitol., 56: 1013–1022.

Hoogstraal, H. 1980. Ticks (Ixodoidea) from Oman. J. Oman Stud. Spec. Rep. (2): 265–272.

Hoogstraal, H. & Kim, K.C. 1985. Tick and mammal coevolution, with emphasis on Haemaphysalis. In K.C. Kim (editor), Coevolution of parasitic arthropods and mammals. John Wiley & Sons, New York, pp. 505–568.

Kaul, H.N., Mishra, A.C., Dhanda, V., Kulkarni, S.M. & Guttikar, S.N. 1978. Ectoparasitic arthropods of birds and mammals from Rajasthan State, India. Ind. J. Parasitol., 2: 19–25.

Keirans, J.E. 1985. George Henry Falkiner Nuttall and the Nuttall tick catalogue. U. S. Dept. Agric., Agric. Res. Ser. Misc. Pub. (1438), 1785 pp.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Mitchell, R.M. & Dick, J.A. 1978. Ectoparasites from Nepal birds. J. Bombay Nat. Hist. Soc., 74: 264–274.

66 – H. indoflava Dhanda & Bhat, 1968 (J. Parasitol., 54: 1063–1067)

Type depositories: VRC (holotype, paratypes), BMNH, HH, IM, USNTC (paratypes) (Keirans and Hillyard 2001, op. cit. under H. aciculifer). The specimens used to describe this species were originally classified as H. flava by Sharif, M. (1928. A revision of the Indian Ixodidae with special reference to the collection in the Indian Museum. Rec. Ind. Mus., 30: 217–344).

Known stages: male, female, nymph, larva

Zoogeographic Region: Oriental

Ecoregions: desert and xeric shrublands

Hosts: Artiodactyla: Bovidae, Suidae; Carnivora: Canidae (A)

Human infestation: yes (Dhanda and Bhat 1968)

Remarks: Camicas et al. (1998) list the larva and nymph of H. indoflava as undescribed, but both stages were earlier described by Dhanda and Bhat (1970), although their hosts remain unknown.

References

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. 1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.

Dhanda, V. & Bhat, H.R. 1968. Haemaphysalis (Haemaphysalis) indoflava sp. n. (Acarina: Ixodidae) from the Himalayan region of Uttar Pradesh, and Madras, India. J. Parasitol., 54: 1063–1067.

Dhanda, V. & Bhat, H.R. 1970. Immature stages of Haemaphysalis indoflava Dhanda and Bhat, 1968 (Acarina: Ixodidae), and notes on its ecology. Oriental Insects, 4: 253–258.

67 – H. inermis Birula, 1895 (Izv. Imp. Akad. Nauk, Ser. 5, 2 (4): 353–364)

Type depository: ZIAC (holotype) (Filippova 2008, op. cit. under H. caucasica). Keirans and Hillyard (2001, op. cit. under H. aciculifer) found that a female of H. inermis sent by Birula to BMNH is possibly the holotype, but the information in Filippova (2008, op. cit. under H. caucasica) contradicts this opinion. See “remarks on some invalid names” for a new synonym of H. inermis.

Known stages: male, female, nymph, larva

Zoogeographic Region: Palearctic

Ecoregions: temperate broadleaf and mixed forests

Hosts: Aves, Squamata and Testudines are considered exceptional hosts for this tick.

Artiodactyla: Suidae (ANL)

Mammalia (several orders) (AN)

Rodentia: Cricetidae, Muridae; Squamata: Lacertidae, Viperidae; Testudines: Testudinidae (NL)

Passeriformes: Sylviidae, Muscicapidae (L)

Charadriiformes: Scolopacidae; Galliformes: Phasianidae; Passeriformes: Corvidae (stages unknown)

Human infestation: yes (Černý 1972; Bursali et al. 2012 among others)

Remarks: there are several records of H. inermis for Taiwan (Oriental Region); however, Robbins (2005) considers them unsound and we concur. Kolonin (2009) excludes Aves, Squamata and Testudinidae as hosts of H. inermis, but they are listed as hosts of this species in Hoogstraal et al. (1964), Nosek et al. (1967), Filippova (1997) and Široký et al. (2006). Nevertheless, records from these hosts appear to be exceptional.

References

Bursali, A., Keskin, A. & Tekin, S. 2012. A review of the ticks (Acari: Ixodida) of Turkey: species diversity, hosts and geographical distribution. Exp. Appl. Acarol., 57: 91–104.

Černý, V. 1972. The tick fauna of Czechoslovakia. Folia Parasitol., 19: 87–92.

Filippova, N.A. 1997. Ixodid ticks of subfamily Amblyomminae. Fauna of Russia and neighbouring countries, 4 (5), Nauka, St. Petersburg, 436 pp. In Russian.

Gusev, V.M., Bednyy, S.N., Guseva, A.A., Labunets, N.F. & Bakeyev, N.N. 1961. The Ecological groups of birds on the Caucasus and their role in the life cycle of ticks and fleas. Trudy Nauch.-Issled. Protiv. Inst. Kavk. Zakav. (5): 217–267. In Russian.

Hoogstraal, H., Traylor, M.A., Gaber, S., Malakatis, G., Guindy, E. & Helmy, I. 1964. Ticks (Ixodidae) on migrating birds in Egypt, spring and fall 1962. Bull. World Health Org., 30: 355–367.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Nosek, J., Lichard, M. & Sztankay, M. 1967. The ecology of ticks in the Tribeč and Hronský Inovec Mountains. Bull. World Health Org., 36 (Suppl. 1): 49–59.

Robbins, R.G. 2005. The ticks (Acari: Ixodida: Argasidae, Ixodidae) of Taiwan: a synonymic checklist. Proc. Entomol. Soc. Wash., 107: 245–253.

Široký, P., Petrželková, K.J., Kamler, D., Mihalca, A.D. & Modrý, D. 2006. Hyalomma aegyptium as dominant tick in tortoises of the genus Testudo in Balkan countries, with notes on its host preferences. Exp. Appl. Acarol., 40: 279–290.

68 – H. intermedia Nuttall & Warburton, 1909 (Parasitology, 2: 57–76)

See H. parva.

Type depository: BMNH (lectotype) (Keirans and Hillyard 2001, op. cit. under H. aciculifer) originally named Haemaphysalis bispinosa intermedia

Known stages: male, female, nymph, larva

Zoogeographic Region: Oriental

Ecoregions: tropical and subtropical dry broadleaf forests; few ticks in desert and xeric shrublands

Hosts: usual hosts for adult ticks are Artiodactyla: Bovidae.

Mammalia (several orders); Cuculiformes: Cuculidae; Galliformes: Phasianidae (ANL)

Passeriformes: Muscicapidae (AN)

Passeriformes (several families) (NL)

Piciformes: Picidae; Columbiformes: Columbidae (L)

Human infestation: no

References

Bhat, H.R. & Sreenivasan, M.A. 1981. Further records of the ticks of some reptilian and mammalian hosts in the Kyasanur Forest disease area, Karnataka, India. Ind. J. Parasitol., 5: 207–210.

Bhat, H.R., Sreenivasan, M.A. & Jacob, P.G. 1986. Ixodid ticks infesting goats in the Kyasanur Forest disease area, Shimoga District, Karnataka. Ind. J. Parasitol., 10: 39–45.

Geevarghese, G. & Dhanda, V. 1995. Ixodid ticks of Maharashtra State, India. Acarologia, 36: 309–313.

Rajagopalan, P.K. 1972. Ixodid ticks (Acarina: Ixodidae) parasitizing wild birds in the Kyasanur Forest disease area of Shimoga District, Mysore State, India. J. Bombay Nat. Hist. Soc., 69: 55–78.

Rebello, M.J. & Reuben, R. 1967. A report on ticks collected from birds and small mammals in North Arcot and Chittoor Districts, South India. J. Bombay Nat. Hist. Soc., 68: 283–289.

Seneviratna, P. 1965. The Ixodoidea (ticks) of Ceylon. Parts II and III. Ceylon Vet. J., 13: 28–54.

69 – H. japonica Warburton, 1908 (Proc. Cambr. Phil. Soc., 14: 508–519)

It is generally held that this taxon comprises two subspecies: H. japonica japonica Warburton, 1908, and H. japonica douglasi Nuttall & Warburton, 1915. However, Japanese workers such as Kitaoka, S. (1985. Keys to the species immature stages of the Japanese Haemaphysalis ticks (Ixodidae). Bull. Natl. Inst. Anim. Health Q., 88: 49–63. In Japanese) and Fujita, H. & Takada, N. (2007. Identification of immature ticks in Japan. In Organization Committee of SADI (editor), Acari and Emerging/Reemerging Diseases. Zenkoku Nosom Kyoiku Kyokai Publishing Company, Tokyo, pp. 53–68. In Japanese, Spanish translation of legends of figures by Kori, V.), among others, treat H. douglasi and H. japonica as distinct species. Filippova, N.A. (1997. Ixodid ticks of subfamily Amblyomminae. Fauna of Russia and neighbouring countries, 4 (5), Nauka, St. Petersburg, 436 pp. In Russian) considers H. douglasi a subspecies of H. japonica, while Yamaguti et al. (1971, op. cit. under H. ias) list only H. japonica as valid, but stress that additional studies are needed to elucidate the relationship between H. japonica and H. japonica douglasi.

Type depository: BMNH (lectotype, paralectotypes) (Keirans and Hillyard 2001, op. cit. under H. aciculifer) as Haemaphysalis japonnica (lapsus).

Known stages: male, female, nymph, larva

Zoogeographic Region: Palearctic

Ecoregions: temperate broadleaf and mixed forests

Hosts: Artiodactyla: Cervidae; Carnivora: Canidae, Mustelidae; Galliformes: Phasianidae; Passeriformes (several families) (ANL)

Artiodactyla: Bovidae; Carnivora: Felidae, Ursidae; Lagomorpha: Leporidae (AN)

Artiodactyla: Suidae; Erinaceomorpha: Erinaceidae; Perissodactyla: Equidae (A)

Rodentia: Cricetidae, Muridae, Sciuridae; Strigiformes: Strigidae (NL)

Charadriiformes: Scolopacidae (N)

Human infestation: yes (Yamauchi et al. 2010)

Remarks: Camicas et al. (1998) do not regard Aves as important hosts for this tick species, but the studies of Emel’yanova and Goordeeva (1969) and others indicate that the opposite is true, especially for larvae and nymphs of H. japonica. Kitaoka (1985) provides a record of H. japonica (named H. douglasi) from Ursidae, but we were unable to determine whether his specimen is a nymph or a larva. See above.

References

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. 1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.

Chen, Z., Yang, X., Bu, F., Yang, X., Yang, X. & Liu, J. 2010. Ticks (Acari: Ixodoidea: Argasidae, Ixodidae) of China. Exp. Appl. Acarol., 51: 393–404.

Emel’yanova, N.D. & Gordeeva, V.P. 1969. Data on the study of birds acting as hosts of ixodid ticks in East Manchurian mountain area spurs and in the Pre-Khanka Plain. In A.O. Cherepanov (editor), Migratory birds and their role in arboviral distribution. Sibirisk. Otd. Akad. Nauk SSSR, Biol. Inst., Akad. Nauk SSSR, Inst. Polio. Virus Entsef., Min. Zdravookhr. RSFSR, Omsk Inst. Prirod.-Ochag. Infekts., Novosibirsk, pp. 160–169. In Russian, NAMRU-3 translation 621.

Filippova, N.A. 1997. Ixodid ticks of subfamily Amblyomminae. Fauna of Russia and neighbouring countries, 4 (5), Nauka, St. Petersburg, 436 pp. In Russian.

Kislenko, G.S. & Koneva, I.V. 1965. Infestations of birds by ixodid ticks in secondary forests in lower reaches of the Ussuri River. Mater. 4. Vses. Orinthol. Konf., Alma-Ata, Sep. 1965, pp. 160–163. In Russian, NAMRU-3 translation 408.

Kitaoka, S. 1985. Keys to the species in immature stages of the Japanese Haemaphysalis ticks (Ixodidae). Bull. Natl. Inst. Anim. Health Q. (88): 49–63. In Japanese.

Shimada, Y., Inokuma, H., Beppu, T., Okuda, M. & Onishi, T. 2003. Survey of ixodid tick species on domestic cats in Japan. Vet. Parasitol., 111: 231–239.

Teng, K.-F. & Jiang, Z.-J. 1991. Economic insect fauna of China. Fasc. 39, Acari: Ixodidae. Science Press, Beijing, 355 pp. In Chinese.

Voloshina, I.V. 2010. Ticks (Ixodidae) of the Lazovsky Reserve. Proc. Sci. Conf. devoted 75 Anniv. Lazovsky Reserve, Vladivostok, pp. 59–65. In Russian.

Yamaguti, N., Tipton, V.J., Keegan, H.L. & Toshioka, S. 1971. Ticks of Japan, Korea and the Ryukyu Islands. Brigham Young Univ. Sci. Bull. Biol. Ser., 15 (1), 226 pp.

Yamauchi, T., Fukui, Y., Watanabe, M., Nakagawa, H. & Kamimura, K. 2010. Forty cases of human infestations with hard ticks (Acari: Ixodidae) in Toyama Prefecture, Japan. Med. Entomol. Zool., 61: 133–143. In Japanese.

70 – H. juxtakochi Cooley, 1946 (Natl. Inst. Health Bull. (187), 54 pp)

Rawlins, S.C., Mahadeo, S. & Martínez, R. (1993. A list of the ticks affecting man and animals in the Caribbean. CARAPHIN News, (6): 8–9) refer to this species as H. juxtakichi and treat its synonym H. kochi Aragão, 1908 as a valid species. See remarks on invalid names at the beginning of this chapter for the synonym of H. kochi Aragão, 1908 and H. juxtakochi.

Type depositories: USNTC (holotype, paratypes), MCZ, DEEZ (paratypes) (Cooley, 1946, op. cit. under H. chordeilis)

Known stages: male, female, nymph, larva

Zoogeographic Regions: Nearctic, Neotropical

Ecoregions: several different Nearctic and Neotropical ecoregions

Hosts: usual hosts for adult ticks are Artiodactyla: Cervidae.

Mammalia (several orders) (ANL)

Passeriformes: Thraupidae (A)

Passeriformes: Corvidae, Thamnophilidae, Turdidae (NL)

Falconiformes: Accipitridae (N)

Galliformes: Cracidae; Passeriformes: Emberizidae (L)

Human infestation: yes (Bermúdez et al. 2012)

Remarks: Camicas et al. (1998) state that this species is found only in the Neotropical Region, but field records from Tamaulipas (Mexico) and Ohio (USA) in Kohls (1960) and Keirans and Restifo (1993) belong to the Nearctic Region. Hoogstraal and Kim (1985) assert that immature stages of H. juxtakochi are primarily parasites of Cervidae and Rodentia: Dasyproctidae; however, the literature does not yield data confirming this argument.

References

Arzua, M., Onofrio, V.C. & Barros-Battesti, D.M. 2005. Catalogue of the tick collection (Acari, Ixodida) of the Museu de História Natural Capão da Imbuia, Curitiba, Paraná, Brazil. Rev. Bras. Zool., 22: 623–632.

Beldoménico, P.M., Baldi, J.C., Antoniazzi, L.R., Orduna, G.M., Mastropaolo, M., Macedo, A.C., Ruiz, M.F., Orcellet, V., Peralta, J.L., Venzal, J.M., Mangold, A.J. & Guglielmone, A.A. 2003. Ixodid ticks (Acari: Ixodidae) present at Parque Nacional El Rey, Argentina. Neotr. Entomol., 32: 273–277.

Bermúdez, S.E., Castro, A., Esser, H., Liefting, Y., García, G. & Miranda, R.J. 2012. Ticks (Ixodida) on humans from central Panama, Panama (2010–2011). Exp. Appl. Acarol., 58: 81–88.

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. 1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.

Debárbora, V.N., Nava, S., Cirignoli, S., Guglielmone, A.A. & Poi, A.S.G. 2012. Ticks (Acari: Ixodidae) parasitizing endemic and exotic wild mammals in the Esteros del Iberá wetlands, Argentina. Syst. Appl. Acarol., 17: 243–250.

Fairchild, G.B., Kohls, G.M. & Tipton, V.J. 1966. The ticks of Panama (Acarina: Ixodoidea). In W.R. Wenzel & V.J. Tipton (editors), Ectoparasites of Panama. Field Museum of Natural History, Chicago, pp. 167–219.

Guglielmone, A.A., Mangold, A.J. & Aufranc, C.R. 1992. Haemaphysalis juxtakochi, Ixodes pararicinus (Ixodidae) and Otobius megnini (Argasidae) in relation to the phytogeography of Argentina. Ann. Parasitol. Hum. Comp., 67: 91–93.

Guglielmone, A.A., Estrada-Peña, A., Keirans, J.E. & Robbins, R.G. 2003. Ticks (Acari: Ixodida) of the Neotropical Zoogeographic Region. Special Publication of the International Consortium on Ticks and Tick-borne Diseases-2, Atalanta, Houten, The Netherlands, 173 pp.

Guglielmone, A.A., Romero, J., Venzal, J.M., Nava, S., Mangold, A.J. & Villavicencio, J. 2005. First record of Haemaphysalis juxtakochi Cooley, 1946 (Acari: Ixodidae) from Peru. Syst. Appl. Acarol., 10: 33–35.

Hoogstraal, H. & Kim, K.C. 1985. Tick and mammal coevolution, with emphasis on Haemaphysalis. In K.C. Kim (editor), Coevolution of parasitic arthropods and mammals. John Wiley & Sons, New York, pp. 505–568.

Ivancovich, J.C. & Luciani, C.A. 1992. Las garrapatas de Argentina. Monografía de la Asociación Argentina de Parasitología Veterinaria, Buenos Aires, 95 pp.

Jones, E.K., Clifford, C.M., Keirans, J.E. & Kohls, G.M. 1972. The ticks of Venezuela (Acarina: Ixodoidea) with a key to the species of Amblyomma in the Western Hemisphere. Brigham Young Univ. Sci. Bull. Biol. Ser., 17 (4), 40 pp.

Keirans, J.E. & Restifo, R.A. 1993. Haemaphysalis juxtakochi Cooley (Acari: Ixodidae), a Neotropical tick species, found in Ohio. J. Med. Entomol., 30: 1074–1075.

Kohls, G.M. 1960. Records and new synonymy of New World Haemaphysalis ticks, with descriptions of the nymph and larva of H. juxtakochi Cooley. J. Parasitol., 46: 355–361.

Labruna, M.B., Barbieri, F.S., Martins, T.F., Brito, L.G. & Ribeiro, F.D.S. 2010. New tick records in Rondônia, western Brazilian Amazon. Rev. Bras. Parasitol. Vet., 19 (3): 192–194.

Nava, S., Lareschi,M., Rebollo, C., Benítez Usher, C., Beati, L., Robbins, R.G., Durden, L.A., Mangold, A.J. & Guglielmone, A.A. 2007. The ticks (Acari: Ixodida: Argasidae, Ixodidae) of Paraguay. Ann. Trop. Med. Parasitol., 101: 255–270.

Ogrzewalska, M., Uezu, A. & Labruna, M.B. 2010. Ticks (Acari: Ixodidae) infesting wild birds in the eastern Amazon, northern Brazil, with notes on rickettsial infection in ticks. Parasitol. Res., 106: 809–816.

Ortiz, F., Nava, S. & Guglielmone, A.A. 2011. Análisis de una colección de garrapatas (Acari: Argasidae, Ixodidae) del norte argentino. Rev. FAVE Cienc. Vet., 10: 49–55.

Venzal, J.M., Félix, M.L., Olmos, A., Mangold, A.J. & Guglielmone, A.A. 2003. A collection of ticks (Ixodidae) from wild birds in Uruguay. Exp. Appl. Acarol., 36: 325–331.

71 – H. kadarsani Hoogstraal & Wassef, 1977 (J. Parasitol., 63: 1103–1109)

Type depositories: MZB (holotype, paratypes) USNTC (paratypes) (Keirans and Hillyard 2001, op. cit. under H. aciculifer) state that there are no paratypes of H. kadarsani in BMNH, despite the assertion by the authors of this species.

Known stages: male, female, nymph

Zoogeographic Region: Australasian

Ecoregion: Sulawesi montane rain forests

Hosts: Rodentia: Muridae (AN)

Human infestation: no

Remarks: Camicas et al. (1998) state that H. kadarsani is found in the Oriental Zoogeographic Region, but this tick is found exclusively on Sulawesi Island, which lies within the Australasian Zoogeographic Region. Durden and Keirans (1996) regard H. kadarsani as an endangered species.

References

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. 1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.

Durden, L.A. & Keirans, J.E. 1996. Host-parasite coextinction and the plight of tick conservation. Am. Entomol., 42: 87–91.

Durden, L.A., Merker, S. & Beati, L. 2008. The tick fauna of Sulawesi, Indonesia (Acari: Ixodoidea: Argasidae and Ixodidae). Exp. App. Acarol., 45: 85–110.

Hoogstraal, H. & Wassef, H.Y. 1977. Haemaphysalis (Ornithophysalis) kadarsani sp. n. (Ixodoidea: Ixodidae), a rodent parasite of virgin lowland forests in Sulawesi (Celebes). J. Parasitol., 63: 1103–1109.

72 – H. kashmirensis Hoogstraal & Varma, 1962 (J. Parasitol., 48: 185–194)

Type depositories: USNTC (holotype, paratypes), VRC, HH, ZSI (paratypes) (Hoogstraal, H. & Varma, M.G.R. 1962. Haemaphysalis cornupunctata sp. n. and H. kashmirensis sp. n. from Kashmir, with notes on H. sundrai Sharif and H. sewelli Sharif of India and Pakistan (Ixodoidea: Ixodidae). J. Parasitol., 48: 185–194)

Known stages: male, female, nymph, larva

Zoogeographic Region: Oriental

Ecoregion: northwestern thorn scrub forests

Hosts: usual hosts for adult ticks are Artiodactyla: Bovidae; usual hosts for larvae and nymphs are Squamata: Agamidae. Squamata are considered exceptional hosts for adults of this tick.

Squamata: Agamidae (ANL)

Artiodactyla: Bovidae (A)

Rodentia: Muridae, Sciuridae (N)

Human infestation: no

Remarks: a record of adult H. kashmirensis from Agamidae in Hoogstraal and Kim (1985) is ignored in Kolonin (2009).

References

Hoogstraal, H. & Kim, K.C. 1985. Tick and mammal coevolution, with emphasis on Haemaphysalis. In K.C. Kim (editor), Coevolution of parasitic arthropods and mammals. John Wiley & Sons, New York, pp. 505–568.

Hoogstraal, H. & McCarthy, V.C. 1965. Hosts and distribution of Haemaphysalis kashmirensis with descriptions of immature stages and definition of the subgenus Herpetobia Canestrini (resurrected). J. Parasitol., 51: 674–679.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

73 – H. kinneari Warburton, 1913 (Parasitology, 6: 121–130)

See H. toxopei.

Type depository: BMNH (holotype) (Keirans and Hillyard 2001, op. cit. under H. aciculifer)

Known stages: male, female, nymph, larva

Zoogeographic Region: Oriental

Ecoregion: Deccan thorn scrub forests

Hosts: Carnivora: Canidae; Lagomorpha: Leporidae (ANL)

Carnivora: Herpestidae (AN)

Carnivora: Felidae, Ursidae (AL)

Artiodactyla: Suidae (A)

Mammalia (several orders); Galliformes: Phasianidae; Cuculiformes: Cuculidae; Passeriformes: Timaliidae, Sylviidae (NL)

Rodentia: Hystricidae; Coraciiformes: Bucerotidae; Passeriformes (several families); Piciformes: Capitonidae (L)

Human infestation: no

Remarks: Camicas et al. (1998) list the larva of H. kinneari as undescribed, but it had earlier been described by Trapido et al. (1964). With the exception of Bhat and Sreenivasan (1981), Rahman and Mondal (1985) and Geevarghese et al. (1997), all authors refer to this species as H. papuana kinneari. Geevarghese et al. (1997) limit the host range of H. kinneari to “tiger, wild boar, monkey and rodent” without elaboration.

References

Bhat, H.R. & Sreenivasan, M.A. 1981. Further records of the ticks of some reptilian and mammalian hosts in the Kyasanur Forest disease area, Karnataka, India. Ind. J. Parasitol., 5: 207–210.

Bhat, H.R., Sreenivasan, M.A. & Jacob, P.G. 1986. Ixodid ticks infesting goats in the Kyasanur Forest disease area, Shimoga District, Karnataka. Ind. J. Parasitol., 10: 39–45.

Boshell, J. & Rajagopalan, P.K. 1968. Observations on the experimental exposure of monkeys, rodents and shrews to infestation of ticks in forest in Kyasanur Forest disease area. Ind. J. Med. Res., 56 (4) (Suppl.): 573–588.

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. 1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.

Geevarghese, G., Fernandes, S. & Kulkarni, S.M. 1997. A checklist of Indian ticks (Acari: Ixodoidea). Ind. J. Anim. Sci., 67: 566–574.

Rahman, M.H. & Mondal, M.M.H. 1985. Tick fauna of Bangladesh. Ind. J. Parasitol., 9: 145–149.

Rajagopalan, P.K. 1972. Ixodid ticks (Acarina: Ixodidae) parasitizing wild birds in the Kyasanur Forest disease area of Shimoga District, Mysore State, India. J. Bombay Nat. Hist. Soc., 69: 55–78.

Rajagopalan, P.K. & Sreenivasan, M.A. 1981. Ixodid ticks on cattle and buffaloes in the Kyasanur Forest disease area of Karnataka State. Ind. J. Med. Res., 73: 880–889.

Rajagopalan, P.K., Patil, A.P. & Boshell, J. 1968. Ixodid ticks on their mammalian hosts in the Kyasanur Forest disease area of Mysore State, India, 1961–1964. Ind. J. Med. Res., 56: 510–526.

Trapido, H., Hoogstraal, H. & Varma, M.G.R. 1964. Status and descriptions of Haemaphysalis p. papuana Thorell (n. comb.) and of H. papuana kinneari Warburton (n. comb.) (Ixodoidea, Ixodidae) of southern Asia and New Guinea. J. Parasitol., 50: 172–188.

74 – H. kitaokai Hoogstraal, 1969 (J. Parasitol., 55: 211–221)

Type depositories: USNTC (holotype, paratypes), NIAH (paratypes) (Hoogstraal, H. 1969. Haemaphysalis (Alloceraea) kitaokai sp. n. of Japan, and keys to species in the structurally primitive subgenus Alloceraea Schulze of Eurasia (Ixodoidea, Ixodidae). J. Parasitol., 55: 211–221)

Known stages: male, female, nymph, larva

Zoogeographic Regions: Oriental, Palearctic

Ecoregions: temperate conifer and broadleaf forests

Hosts: Aves are considered exceptional hosts for this tick.

Artiodactyla: Bovidae, Cervidae (AN)

Perissodactyla: Equidae; Galliformes: Phasianidae (A)

Human infestation: yes (Mahara 1997)

Remarks: the larva of H. kitaokai is known only from laboratory-reared specimens (Hoogstraal 1969). Camicas et al. (1998) state that this species is exclusively Palearctic, but Teng and Jiang (1991) show that the distribution of H. kitaokai encompasses localities in the Oriental and Palearctic Zoogeographic Regions. Xu and Li (1997) found a female of H. kitaokai on Phasianidae, but Kolonin (2009) ignores Aves as hosts for adult ticks.

References

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. 1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.

Chen, Z., Yang, X., Bu, F., Yang, X., Yang, X. & Liu, J. 2010. Ticks (Acari: Ixodoidea: Argasidae, Ixodidae) of China. Exp. Appl. Acarol., 51: 393–404.

Hoogstraal, H. 1969. Haemaphysalis (Alloceraea) kitaokai sp. n. of Japan, and keys to species in the structurally primitive subgenus Alloceraea Schulze of Eurasia (Ixodoidea, Ixodidae). J. Parasitol., 55: 211–221.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Mahara, F. 1997. Japanese spotted fever: report of 31 cases and review of the literature. Emerg. Inf. Dis., 3: 105–111.

Teng, K.-F. & Jiang, Z.-J. 1991. Economic insect fauna of China. Fasc. 39, Acari: Ixodidae. Science Press, Beijing, 355 pp. In Chinese.

Xu, R. & Li, K. 1997. A collection of ticks from Guizhou, China. Syst. Appl. Acarol., 2: 245–246.

Xu, R. & Luo, G. 1998. Ticks parasitic on Muntiacus reevesi and Lepus sinensis in Wuyi Mountains, Fujian, China. Syst. Appl. Acarol. 3: 197.

Yamaguti, N., Tipton, V.J., Keegan, H.L. & Toshioka, S. 1971. Ticks of Japan, Korea and the Ryukyu Islands. Brigham Young Univ. Sci. Bull. Biol. Ser., 15 (1), 226 pp.

Yamauchi, T. 2005. A bibliographical survey on ixodid fauna of Shimane Prefecture, Japan (Acari: Ixodidae). Bull. Hoshizaki Green Found., 8: 289–301. In Japanese.

75 – H. knobigera Prakasan & Ramani, 2007 (Int. J. Zool. Res., 3: 169–177)

This species is not included in Kolonin (2009, op. cit. under H. anomaloceraea). Guglielmone et al. (2009, op. cit. under H. colasbelcouri) doubt the validity of this species, whose description is very poor, but have included it in their list of valid names because it is uncertain whether it is a synonym of another species of Haemaphysalis found on Bovidae in India (Oriental Region).

Type depository: DA (holotype, paratypes) (Prakasan, K. & Ramani, M. 2007. Two new species of ixodid ticks (Acarina: Ixodida) from Kerala, India. Int. J. Zool. Res., 3: 169–177)

Known stages: male

Zoogeographic Region: Oriental

Ecoregion: southwestern Ghats moist deciduous forests

Hosts: Artiodactyla: Bovidae (A)

Human infestation: no

Reference

Prakasan, K. & Ramani, M. 2007. Two new species of ixodid ticks (Acarina: Ixodida) from Kerala, India. Int. J. Zool. Res., 3: 169–177.

76 – H. koningsbergeri Warburton & Nuttall, 1909 (Parasitology, 2: 57–76)

Type depository: BMNH (lectotype, paralectotypes) (Keirans and Hillyard 2001, op. cit. under H. aciculifer)

Known stages: male, female, larva

Zoogeographic Region: Oriental

Ecoregions: tropical and subtropical moist broadleaf forests

Hosts: usual hosts for adult ticks are Carnivora (several families).

Carnivora: Mustelidae; Rodentia: Muridae (A, N and/or L)

Mammalia (several orders) (A)

Human infestation: yes (Keirans 1985)

Remarks: Camicas et al. (1998) state that the larva of H. koningsbergeri is undescribed, but this stage was described by Kadarsan (1971). Tanskul et al. (1983) use the term “immatures” without stating whether larvae, nymphs or both stages were found on hosts, but we consider their records provisionally valid. There is a valid record of H. koningsbergeri from a human in Audy et al. (1960), but the specimen was merely crawling on the body and we therefore exclude humans as hosts of this tick. It is uncertain whether the nymph found in a nest of Pilocercidae by Kohls (1957) actually fed on this type of host, and Hoogstraal et al. (1972) treat the determination of a nymph of H. koningsbergeri on Sciuridae as tentative, while regarding a collection from Phasianidae (tick stage unknown) as probably erroneous. These records have not been included in our host list for this species.

References

Audy, J.R., Nadchatram, M. & Lim, B.-L. 1960. Malaysian parasites. XLIX. Host distribution of Malayan ticks (Ixodoidea). Stud. Inst. Med. Res. Malaya, 29: 225–246.

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. 1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.

Hoogstraal, H., Lim, B.L., Nadchatram, M. & Anastos, G. 1972. The Gunong Benom Expedition 1967. 8. Ticks (Ixodidae) of Gunong Benom and their altitudinal distribution, hosts and medical relationships. Bull. Br. Mus. (Nat. Hist.) Zool., 23: 167–186.

Kadarsan, S. 1971. Larval ixodid ticks of Indonesia (Acarina: Ixodidae). Ph.D. Dissertation, University of Maryland, 182 pp.

Keirans, J.E. 1985. George Henry Falkiner Nuttall and the Nuttall tick catalogue. U. S. Dept. Agric., Agric. Res. Ser. Misc. Pub. (1438), 1785 pp.

Kohls, G.M. 1957. Malaysian parasites XVIII. Ticks (Ixodoidea) of Borneo and Malaya. Malaya (28): 65–94.

Tanskul, P., Stark, H.E. & Inlao, I. 1983. A checklist of ticks of Thailand (Acari: Metastigmata: Ixodoidea). J. Med. Entomol., 20: 330–341.

77 – H. kopetdaghica Kerbabaev, 1962 (Akad. Nauk. Turkmenistan SSR Ser. Biol. Nauk (1): 77–80. In Russian)

Type depository: ZIAC (neotype) (Filippova 2008, op. cit. under H. caucasica), originally named H. warburtoni kopetdaghicus.

Known stages: male, female, nymph, larva

Zoogeographic Region: Palearctic

Ecoregion: Kopet Dag semi-desert

Hosts: usual hosts for larvae, nymphs and adult ticks are Artiodactyla: Bovidae.

Artiodactyla: Bovidae (ANL)

Perissodactyla: Equidae (AN)

Carnivora: Felidae (A)

Human infestation: no

Remarks: Mihalca et al. (2011) regard this species as endangered.

References

Filippova, N.A. 1997. Ixodid ticks of subfamily Amblyomminae. Fauna of Russia and neighbouring countries, 4 (5), Nauka, St. Petersburg, 436 pp. In Russian.

Hoogstraal, H. & Wassef, H.Y. 1979. Haemaphysalis (Allophysalis) kopetdaghica: identity and discovery of each feeding stage on the wild goat in northern Iran (Ixodoidea: Ixodidae). J. Parasitol., 65: 783–790.

Mihalca, A.D., Gherman, C.M. & Cozma, V. 2011. Coendangered hard-ticks: threatened or threatening? Parasit. Vectors, 4 (71), 7 pp.

78 – H. kumaonensis Geevarghese & Mishra, 2011 (In Geevarghese, G. & Mishra, A.C. 2011. Haemaphysalis ticks of India. Elsevier, London, 260 pp.)

Type depositories: not stated in Geevarghese, G. & Mishra, A.C. (2011. Haemaphysalis ticks of India. Elsevier, London, 260 pp.)

Known stages: male, female, nymph, larva

Zoogeographic Region: undetermined

Ecoregions: undetermined

Hosts: unknown

Human infestation: no

Remarks: no locality or host data are provided for this species, and its relationship to other taxa is uncertain from the description in Geevarghese and Mishra (2011). We therefore consider this species provisionally valid.

Reference

Geevarghese, G. & Mishra, A.C. 2011. Haemaphysalis ticks of India. Elsevier, Amsterdam and several other cities, 260 pp.

79 – H. kutchensis Hoogstraal & Trapido, 1963 (J. Parasitol., 49: 489–497)

Type depositories: USNTC (holotype, paratypes), BMNH, ZIAC, FMNH, HH, VRC (paratypes) (Keirans and Hillyard 2001, op. cit. under H. aciculifer). This species was originally identified as H. bispinosa intermedia Warburton and Nuttall, 1909, a synonym of H. intermedia, by Nuttall and Warburton (1915, op. cit. under H. celebensis), as discussed in Hoogstraal, H. & Trapido, H. (1963. Haemaphysalis kutchensis sp. n., a common larval and nymphal parasite of birds in northwestern India (Ixodoidea, Ixodidae). J. Parasitol., 49: 489–497).

Known stages: male, female, nymph, larva

Zoogeographic Regions: Oriental, Palearctic

Ecoregions: desert and xeric shrublands

Hosts: Passeriformes: Sylviidae (ANL)

Carnivora: Canidae (A, N and/or L)

Rodentia: Muridae (A and/or N and/or L)

Artiodactyla: Bovidae; Carnivora: Felidae, Hyaenidae; Lagomorpha: Leporidae; Galliformes: Phasianidae; Cuculiformes: Cuculidae; Strigiformes: Strigidae (A)

Passeriformes (several families) (N)

Human infestation: no

Remarks: Camicas et al. (1998) state that this tick is found only in the Oriental Region, but following Filippova (1997), we consider it to also be present in the Palearctic Region. There is an Afrotropical record in Hoogstraal and Wassef (1985) from a migrant bird, but there is no evidence that H. kutchensis has become established in that region. Kaul et al. (1978) generalized that adults, nymphs and larvae of H. kutchensis are found on Muridae and Canidae without specifying which stages are found on each type of host. Rao et al. (1973) report parasitism of “mongoose” by H. kutchensis without further data, and this record has not been included in our host list for this species.

References

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. 1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.

Filippova, N.A. 1997. Ixodid ticks of subfamily Amblyomminae. Fauna of Russia and neighbouring countries, 4 (5), Nauka, St. Petersburg, 436 pp. In Russian.

Hoogstraal, H. & Trapido, H. 1963. Haemaphysalis kutchensis sp. n., a common larval and nymphal parasite of birds in northwestern India (Ixodoidea, Ixodidae). J. Parasitol., 49: 489–497.

Hoogstraal, H. & Wassef, H.Y. 1985. Haemaphysalis kutchensis, an Indian-Pakistani bird and mammal tick, parasitizing a migrant whitethroat in the Sultanate of Oman. J. Parasitol., 71: 129–130.

Kaul, H.N., Mishra, A.C., Dhanda, V., Kulkarni, S.M. & Guttikar, S.N. 1978. Ectoparasitic arthropods of birds and mammals from Rajasthan State, India. Ind. J. Parasitol., 2: 19–25.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Rao, T.R., Dhanda, V., Bhat, H.R. & Kulkarni, S.M. 1973. A survey of haematophagous arthropods in western Himalayas, Sikkim and Hill Districts of West Bengal. A general account. Ind. J. Med. Res., 61: 1421–1461.

80 – H . kyasanurensis Trapido, Hoogstraal & Rajagopalan, 1964 (J. Parasitol., 50: 295–302)

Type depositories: USNTC (holotype, paratypes), BMNH, VRC, ZIAC, HH (paratypes). (Trapido, H., Hoogstraal, H. & Rajagopalan, P.K. 1964. Haemaphysalis kyasanurensis sp. n., a member of the formosensis group in southern India and Ceylon (Ixodoidea, Ixodidae). J. Parasitol., 50: 295–302). However, Keirans and Hillyard (2001, op. cit., under H. aciculifer) state that no paratypes of H. kyasanurensis are present in BMNH.

Known stages: male, female, nymph, larva

Zoogeographic Region: Oriental

Ecoregion: south Deccan plateau dry deciduous forests

Hosts: usual hosts for larvae, nymphs and adults are Rodentia: Hystricidae.

Rodentia: Hystricidae; Artiodactyla: Bovidae (ANL)

Carnivora: Canidae; Rodentia: Sciuridae (AN)

Artiodactyla: Suidae; Carnivora: Felidae (A)

Primates: Cercopithecidae; Aves (several orders) (NL)

Mammalia (several orders) (L)

Human infestation: no

References

Bhat, H.R. & Sreenivasan, M.A. 1981. Further records of the ticks of some reptilian and mammalian hosts in the Kyasanur Forest disease area, Karnataka, India. Ind. J. Parasitol., 5: 207–210.

Bhat, H.R., Sreenivasan, M.A. & Jacob, P.G. 1986. Ixodid ticks infesting goats in the Kyasanur Forest disease area, Shimoga District, Karnataka. Ind. J. Parasitol., 10: 39–45.

Boshell, J. & Rajagopalan, P.K. 1968. Observations on the experimental exposure of monkeys, rodents and shrews to infestation of ticks in forest in Kyasanur Forest disease area. Ind. J. Med. Res., 56 (4) (Suppl.): 573–588.

Hoogstraal, H. & Rack, G. 1967. Ticks (Ixodidae) collected by Deutsche Indien-Expedition, 1955–1958. J. Med. Entomol., 3: 284–288.

Rajagopalan, P.K. 1972. Ixodid ticks (Acarina: Ixodidae) parasitizing wild birds in the Kyasanur Forest disease area of Shimoga District, Mysore State, India. J. Bombay Nat. Hist. Soc., 69: 55–78.

Rajagopalan, P.K., Patil, A.P. & Boshell, J. 1968. Ixodid ticks on their mammalian hosts in the Kyasanur Forest disease area of Mysore State, India, 1961–1964. Ind. J. Med. Res., 56: 510–526.

81 – H. lagostrophi Roberts, 1963 (Aust. J. Zool., 11: 35–80)

Type depositories: WAM (holotype, paratypes), ANIC (paratypes) (Roberts 1963, op. cit. under H. bremneri; Halliday, B. personal communication to Guglielmone, A.A.)

Known stages: male, female, nymph

Zoogeographic Region: Australasian

Ecoregion: Kimberly tropical savanna

Hosts: usual hosts for adult ticks are Diprotodontia: Macropodidae.

Diprotodontia: Macropodidae (AN)

Peramelemorphia: Peramelidae (A)

Human infestation: no

References

Roberts, F.H.S. 1963. A systematic study of the Australian species of the genus Haemaphysalis Koch (Acarina: Ixodidae). Aust. J. Zool., 11: 35–80.

Roberts, F.H.S. 1970. Australian ticks. CSIRO, Melbourne, 267 pp.

82 – H. lagrangei Larrousse, 1925 (Ann. Parasitol. Hum. Comp., 3: 301–305)

Type depositories: IP (lectotype), CM, BMNH, USNTC (paralectotypes) (Trapido, H. 1965. Notes on critical Asian Haemaphysalis specimens in European museum collections, with designations of lectotypes and a neotype. Proc. Entomol. Soc. Wash., 67: 152–165; Keirans and Clifford 1984, op. cit. under H. bartelsi; Keirans and Hillyard 2001, op. cit. under H. aciculifer).

Known stages: male, female, nymph, larva

Zoogeographic Region: Oriental

Ecoregions: tropical and subtropical moist broadleaf forests

Hosts: usual hosts for adult ticks are Artiodactyla: Cervidae and Carnivora: Mustelidae and Viverridae; usual hosts for larvae and nymphs are Carnivora: Mustelidae and Viverridae. Aves and Squamata are considered exceptional hosts for this tick.

Mammalia (several orders) (ANL)

Galliformes: Phasianidae (AN)

Falconiformes: Falconidae; Passeriformes: Laniidae, Muscicapidae; Squamata: Varanidae (A)

Human infestation: yes (Tanskul et al. 1983)

Remarks: Phan Trong (1977), who refers to this tick as H. hystricis indochinensis Phan Trong, 1977, found this species on Squamata and Aves apart from mammals. Two records of Aves as hosts for H. lagrangei are also included in Hoogstraal et al. (1973). All these records have been ignored in Kolonin (2009), probably because of their infrequency.

References

Chen, Z., Yang, X., Bu, F., Yang, X., Yang, X. & Liu, J. 2010. Ticks (Acari: Ixodoidea: Argasidae, Ixodidae) of China. Exp. Appl. Acarol., 51: 393–404.

Hoogstraal, H., El Kammah, K.M., Santana, F.J. & Van Peenen, P.F.D. 1973. Studies on Southeast Asian Haemaphysalis ticks (Ixodoidea: Ixodidae). H. (Kaiseriana) lagrangei Larrouse: identity, distribution, and hosts. J. Parasitol., 59: 1118–1129.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Phan Trong, C. 1977. Ve bet va con trung ky sinh o Viet Nam. Tap 1. Ve (Ixodoidea), mo ta va phan loai. Ha Noi: Khoa hoc va ky thuat, 489 pp. In Vietnamese.

Tanskul, P., Stark, H.E. & Inlao, I. 1983. A checklist of ticks of Thailand (Acari: Metastigmata: Ixodoidea). J. Med. Entomol., 20: 330–341.

83 – H. laocayensis Phan Trong, 1977 (Ve bet va con trung ky sinh o Viet Nam. Tap 1. Ve (Ixodoidea), mo ta va phan loai. Ha Noi: Khoa hoc va ky thuat, 489 pp.) In Vietnamese.

Type depository: AI (holotype, paratypes) (Kolonin, G.V. personal communication to Guglielmone, A.A.)

Known stages: male, female

Zoogeographic Region: Oriental

Ecoregion: south China-Vietnam subtropical evergreen forests

Hosts: Artiodactyla: Cervidae; Carnivora: Mustelidae (A)

Human infestation: no

Reference

Kolonin, G.V. 1995. Review of the ixodid tick fauna (Acari: Ixodidae) of Vietnam. J. Med. Entomol., 32: 276–282.

84 – H. leachi (Audouin, 1826) (Explication sommaire des planches d’arachnides de l’Égypte et de la Syrie. In Savigny, J. 1826. Description de l’Égypte ou Recueil des Observations et des Recherches qui ont été Faites en Égypte pendant l’Expédition de l’Armée Française. Histoire Naturelle, 1 (4): 99–186. C.L.F. Panckoucke: Paris). Note: major discrepancies exist among citations of this reference.

Type depository: USNTC (neotype) (Keirans and Clifford 1984, op. cit. under H. bartelsi). Originally named Ixodes leachii.

Known stages: male, female, nymph, larva

Zoogeographic Regions: Afrotropical, Palearctic

Ecoregions: several Afrotropical and Palearctic ecoregions

Hosts: usual hosts for adult ticks are Carnivora (several families).

Carnivora (several families); Artiodactyla: Bovidae, Suidae; Primates: Cercopithecidae; Rodentia: Muridae (A)

Human infestation: yes (Apanaskevich et al. 2007)

Remarks: all stages of H. leachi have been misdetermined in the literature, where different species are alleged to constitute the H. leachi group. We have therefore ignored records of this species published prior to Apanaskevich et al. (2007), including Oriental records in Phan Trong (1977). Kolonin (2009) states that the immature stages of this tick feed “on rodents and other small mammals,” supporting the assertion in Apanaskevich et al. (2007). However, the latter authors also state that a taxonomic revision of the H. leachi group will be necessary in order to delimit the host range of the immature stages. For this reason we have not included hosts for larvae and nymphs of this tick. Keirans and Durden (2001) record introductions of H. leachi or ticks from the H. leachi group into the Nearctic Region, but no evidence exists that any member of this group has become established there. See also H. leachi and H. zumpti.

References

Apanaskevich, D.A., Horak, I.G. & Camicas, J.-L. 2007. Redescription of Haemaphysalis (Rhipistoma) elliptica (Koch, 1844), an old taxon of the Haemaphysalis (Rhipistoma) leachi group from East and southern Africa, and of Haemaphysalis (Rhipistoma) leachi (Audouin, 1826) (Ixodida: Ixodidae). Onderstepoort J. Vet. Res., 74: 181–208.

Keirans, J.E. & Durden, L.A. 2001. Invasion: exotic ticks (Acari: Argasidae, Ixodidae) imported into the United States. A review and new records. J. Med. Entomol., 38: 850–861.

Phan Trong, C. 1977. Ve bet va con trung ky sinh o Viet Nam. Tap 1. Ve (Ixodoidea), mo ta va phan loai. Ha Noi: Khoa hoc va ky thuat, 489 pp. In Vietnamese.

85 – H. lemuris Hoogstraal, 1953 (Bull. Mus. Comp. Zool., 111: 37–113)

Type depositories: USNTC (holotype, paratypes), HH, MCZ, OVI (paratypes) (Hoogstraal, H. 1953. Ticks (Ixodoidea) of the Malagasy Faunal Region (excepting the Seychelles). Their origins and host-relationships; with descriptions of five new Haemaphysalis species. Bull. Mus. Comp. Zool., 111: 37–113)

Known stages: male, female, nymph

Zoogeographic Region: Afrotropical

Ecoregion: Madagascar succulent woodlands

Hosts: Primates: Indriidae, Lemuridae (AN)

Primates: Cheirogaleidae, Epilemuridae (N)

Human infestation: no

Remarks: Uilenberg et al. (1979) considered a record of two nymphs of H. lemuris from Passeriformes: Vangidae as uncertain, and this family has been excluded from our host list. Barrett et al. (2012) state that humans, domestic animals and rodents are hosts of H. lemuris, but we were unable to confirm this assertion with the references provided by the authors, and these hosts are not included in the list above. Durden and Keirans (1996) regard H. lemuris as an endangered species.

References

Barrett, M.A., Brown, J.L., Junge, R.E. & Yoder, A.D. 2012. Climate change, predictive modeling and lemur health: assessing impacts of changing climate on health conservation in Madagascar. Biol. Conserv., 157: 409–422.

Durden, L.A. & Keirans, J.E. 1996. Host-parasite coextinction and the plight of tick conservation. Am. Entomol., 42: 87–91.

Durden, L.A., Zohdy, S. & Laakkonen, J. 2010. Lice and ticks of the Eastern rufous mouse lemur, Microcebus rufus, with descriptions of the male and third instar nymph of Lemurpediculus verruculosus (Phthiraptera: Anoplura). J. Parasitol., 96: 874–878.

Uilenberg, G., Hoogstraal, H. & Klein, J.-M. 1979. Les tiques (Ixodoidea) de Madagascar et leur rôle vecteur. Arch. Inst. Pasteur Madagascar Num. Spéc., 153 pp.

86 – H. leporispalustris (Packard, 1869) (First Annual Report of the Trustees of the Peabody Academy of Sciences, Appendix, pp. 56–69)

Type depository: MCZ (type?) (Cooley 1946, op. cit. under H. chordeilis) as Ixodes leporispalustris. Cooley (1946) states that the type of this species is in the MCZ, but on the museum web page for zoological collections (http://mczbase.mcz.harvard.edu/ accessed April 4, 2013), the specimen of Ixodes leporispalustris (as Ixodes leporis-palustris) bears the label “type?”

Known stages: male, female, nymph, larva

Zoogeographic Regions: Nearctic, Neotropical

Ecoregions: several different Nearctic and Neotropical ecoregions

Hosts: usual hosts for larvae, nymphs and adult ticks are Lagomorpha: Leporidae.

Mammalia (several orders); Galliformes: Phasianidae, Odontophoridae; Passeriformes (several families); Strigiformes: Strigidae (ANL)

Cuculiformes: Cuculidae (NL)

Caprimulgiformes: Caprimulgidae; Charadriiformes: Scolopacidae; Columbiformes: Columbidae (N)

Charadriiformes: Laridae, Sternidae; Piciformes: Picidae (L)

Falconiformes: Falconidae; Galliformes: Cracidae (stages unknown)

Human infestation: yes (Furman and Loomis 1984; Guglielmone et al. 2003)

Remarks: Neumann (1897, 1911) states that H. leporispalustris (cited as H. leporis) has been found in the Oriental Region, but we consider this determination erroneous. We also maintain that recent records of H. leporispalustris on Galliformes in Pakistan (Khattak et al. 2012) are in error. More recently, Han et al. (2009) state that H. leporispalustris is found on several wild and domestic animals in South Korea (Palearctic Region); this statement is treated as valid by Sutor et al. (2013), but we consider it to be erroneous. Kolonin (2009) excludes Aves as hosts of adult H. leporispalustris, but we feel that records of this stage from avian hosts listed in Bequaert (1945), Bishopp and Trembley (1945) and Hamer et al. (2012) are valid. Keirans (1985) considers his diagnoses of adults and larvae of H. leporispalustris on Dasyproctidae to be tentative. This mammal has therefore been provisionally excluded from our host list of H. leporispalustris.

References

Bequaert, J.C. 1945. The ticks or Ixodoidea, of the northeastern United States and eastern Canada. Entomol. Am., 25: 73–232.

Bishopp, F.C. & Trembley, H.L. 1945. Distribution and hosts of certain North American ticks. J. Parasitol., 31: 1–54.

Cooley, R.A. 1946. The genera Boophilus, Rhipicephalus, and Haemaphysalis (Ixodidae) of the New World. Natl. Inst. Health Bull. (187), 54 pp.

Furman, D.P. & Loomis, E.C. 1984. The ticks of California (Acari: Ixodida). Bull. Calif. Insect Surv., 25: 1–239.

Galloway, T.D. 2006. Ectoparasites (Phthiraptera: Philopteridae; Acari: Ixodidae) of common nighthawk, Chordeilis minor, and whip-poor-will, Caprimulgus vociferous (Caprimulgiformes: Caprimulgidae), in Manitoba. J. Entomol. Soc. Ontario, 137: 5–11.

Guglielmone, A.A., Estrada-Peña, A., Keirans, J.E. & Robbins, R.G. 2003. Ticks (Acari: Ixodida) of the Neotropical Zoogeographic Region. Special Publication of the International Consortium on Ticks and Tick-borne Diseases-2, Atalanta, Houten, The Netherlands, 173 pp.

Hamer, S.A., Goldberg, T.L., Kitron, U.D., Brawn, J.D., Anderson, T.K., Loss, S.R., Walker, E.D. & Hamer, G.L. 2012. Wild birds and urban ecology of ticks and tick-borne pathogens, Chicago, Illinois, USA, 2005–2010. Emerg. Inf. Dis., 18: 1589–1595.

Han, J.-I., Jang, H.-J. & Na, K.-J. 2009. Molecular detection of Theileria sp. in wild Chinese water deer (Hydropotes inermis argyropus). J. Wildl. Dis., 45: 1213–1216.

Hyland, K.E., Bernier, J., Markowski, D., MacLachlan, A., Amr, Z., Pitocchelli, J., Myers, J. & Hu, R. 2000. Records of ticks (Acari: Ixodidae) parasitizing birds (Aves) in Rhode Island, USA. Int. J. Acarol., 26: 183–192.

Keirans, J.E. 1985. George Henry Falkiner Nuttall and the Nuttall tick catalogue. U. S. Dept. Agric., Agric. Res. Ser. Misc. Pub. (1438), 1785 pp.

Khattak, R.M., Ali, S., Jahangir, M., Khan, M.N., Rasul, A. & Iqbal, F. 2012. Prevalence of ectoparasites in wild and domesticated grey (Francolinus pondicerianus) and black partridge (Francolinus francolinus) from Khyber Pakhtoonkhwa Province of Pakistan. Pakistan J. Zool., 44: 1239–1244.

Kohls, G.M. 1960. Records and new synonymy of New World Haemaphysalis ticks, with descriptions of the nymph and larva of H. juxtakochi Cooley. J. Parasitol., 46: 355–361.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Neumann, L.G. 1897. Révision de la famille des ixodidés (2^e mémoire). Mém. Soc. Zool. Fr., 10: 324–420.

Neumann, L.G. 1911. Ixodidae. Das Tierreich (26), 169 pp.

Samuel, W.M. & Low, W.A. 1970. Parasites of the collared peccary from Texas. J. Wildl. Dis., 6: 16–23.

Scott, J.D., Anderson, J.F. & Durden, L.A. 2012. Widespread dispersal of Borrelia burgdorferi- infected ticks collected from songbirds across Canada. J. Parasitol., 98: 49–59.

Sutor, A., Schwarz, S., & Conraths, F.J. 2013. The biological potential of the raccoon dog (Nyctereutes procyonoides, Gray, 1834) as an invasive species in Europe – new risks for disease spread ? Acta Theriol., doi:10.1007/s13364-013-0138-9

Tugwell, P. & Lancaster, J.L. 1962. Results of a tick-host study in northwest Arkansas. J. Kansas Entomol. Soc., 35: 202–211.

87 – H. lobachovi Kolonin, 1995 (Folia Parasitol., 42: 239)

Type depositories: ZMMO, KC. (Kolonin, G.V. 1995. Haemaphysalis lobachovi sp. n. (Acarina: Ixodidae) from porcupine (Hystrix cristata) from Ethiopia. Folia Parasitol., 42: 239)

Known stages: male

Zoogeographic Region: Afrotropical

Ecoregion: Ethiopian montane moorlands

Hosts: Rodentia: Hystricidae (A)

Human infestation: no

Reference

Kolonin, G.V. 1995. Haemaphysalis lobachovi sp. n. (Acarina: Ixodidae) from porcupine (Hystrix cristata) from Ethiopia. Folia Parasitol., 42: 239.

88 – H. longicornis Neumann, 1901 (Mém. Soc. Zool. Fr., 14: 249–372)

See H. goral and remarks below.

Type depository: ENV (lectotype, paralectotype) (Hoogstraal, H., Roberts, F.H.S., Kohls, G.M. & Tipton, V.J. 1968. Review of Haemaphysalis (Kaiseriana) longicornis Neumann (resurrected) of Australia, New Zealand, New Caledonia, Fiji, Japan, Korea, and northeastern China and USSR, and its parthenogenetic and bisexual populations (Ixodoidea, Ixodidae). J. Parasitol., 54: 1197–1213). See also H. goral.

Known stages: male, female, nymph, larva

Zoogeographic Regions: Australasian, Oriental, Palearctic

Ecoregions: temperate broadleaf and mixed forests

Hosts: usual hosts for adult ticks are Artiodactyla: Bovidae and Cervidae, and Perissodactyla: Equidae. Aves are considered exceptional hosts for this tick.

Mammalia (several orders); Galliformes: Phasianidae (ANL)

Anseriformes. Anatidae (A)

Apterygiformes: Apterygidae; Gruiformes: Rallidae; Passeriformes (several families) (NL)

Psittaciformes: Psittacidae (N)

Charadriiformes: Laridae (L)

Human infestation: yes (Yamauchi et al. 2010)

Remarks: records prior to Hoogstraal et al. (1968) have been ignored here because of their diagnostic uncertainties. A recent study of bisexual and parthenogenetic populations of H. longicornis in China showed that these populations are conspecific (Chen et al. 2012). Hoogstraal et al. (1981) reported the introduction of H. longicornis into the northern sector of the Afrotropical Zoogeographic Region, while Keirans and Durden (2001) reported its introduction into the Nearctic and Hoogstraal et al. (1968) reported its introduction into central Pacific islands, but H. longicornis has apparently failed to become established at any of these localities. According to Heath et al. (1988), infestation of Aves by H. longicornis appears to be an infrequent event, although Hoogstraal et al. (1968) state that Somov and Shestakov (1963), who refer to this tick as H. neumanni, found many birds that were infested with larvae. However, the English translation of the latter study by Russian researchers does not indicate parasitism of Aves by H. longicornis; we therefore accept the view of Heath et al. (1998). See also H. bispinosa and H. davisi.

References

Chen, Z., Yang, X., Bu, F., Ynag, X. & Liu, J. (2012) Morphological and molecular characteristics of bisexual and parthenogenetic Haemaphysalis longicornis. Vet. Parasitol., 189: 344–352.

Clark, J.M., Heath, D.D. & Stankiewicz, M.1997. The ectoparasites of brushtail possum Trichosurus vulpecula in New Zealand. N. Zeal. J. Zool., 24: 199–204.

Filippova, N.A. 1997. Ixodid ticks of subfamily Amblyomminae. Fauna of Russia and neighbouring countries, 4 (5), Nauka, St. Petersburg, 436 pp. In Russian.

Heath, A.C.G. 2013. Implications for New Zealand of potentially invasive ticks sympatric with Haemaphysalis longicornis Neumann, 1901 (Acari: Ixodidae). Syst. Appl. Acarol., 18: 1–26.

Heath, A.C.G., Tenquist, J.D. & Bishop, D.M. 1988. Bird hosts of the New Zealand cattle tick, Haemaphysalis longicornis. N. Zeal. J. Zool., 15: 585–586.

Hoogstraal, H., Roberts, F.H.S., Kohls, G.M. & Tipton, V.J. 1968. Review of Haemaphysalis (Kaiseriana) longicornis Neumann (resurrected) of Australia, New Zealand, New Caledonia, Fiji, Japan, Korea, and northeastern China and USSR, and its parthenogenetic and bisexual populations (Ixodoidea, Ixodidae). J. Parasitol., 54: 1197–1123.

Hoogstraal, H., Wassef, H.Y. & Büttiker, W. 1981. Ticks (Acarina) of Saudi Arabia Fam. Argasidae, Ixodidae. Fauna Saudi Arabia, 3: 25–110.

Kang, J.-G., Kim, H.-C., Choi, C.-Y., Nam, H.-Y., Chae, H.-Y., Chong, S.-T., Klein, T.A., Ko, S. & Chae, J.-S. 2013. Molecular detection of Anaplasma, Bartonella, and Borrelia species in ticks collected from migratory birds from Hong-do Island, Republic of Korea. Vector- Borne Zoon. Dis., 13: 215–225.

Keirans, J.E. 1985. George Henry Falkiner Nuttall and the Nuttall tick catalogue. U. S. Dept. Agric., Agric. Res. Ser. Misc. Pub. (1438), 1785 pp.

Keirans, J.E. & Durden, L.A. 2001. Invasion: exotic ticks (Acari: Argasidae, Ixodidae) imported into the United States. A review and new records. J. Med. Entomol., 38: 850–861.

Roberts, F.H.S. 1970. Australian ticks. CSIRO, Melbourne, 267 pp.

Somov, G.P. & Shestakov, V.I. 1963. On spontaneous infection of Haemaphysalis japonica douglasi Nutt. and Warb. ticks with D. sibiricus rickettsiae in the Primorsk Region. J. Microbiol. (Moscow), 40: 51–56. In Russian, NAMRU-3 translation 205.

Xu, R. & Li, K. 1997. A collection of ticks from Guizhou, China. Syst. Appl. Acarol., 2: 245–246.

Yamauchi, T. 2001. A bibliographical survey of host-parasite relationships between birds and ticks from Japan. Bull. Hoshizaki Green Found., 5: 271–308. In Japanese.

Yamauchi, T., Fukui, Y., Watanabe, M., Nakagawa, H. & Kamimura, K. 2010. Forty cases of human infestations with hard ticks (Acari: Ixodidae) in Toyama Prefecture, Japan. Med. Entomol. Zool., 61: 133–143. In Japanese.

89 – H. luzonensis Hoogstraal & Parrish, 1968 (J. Parasitol., 54: 402–410)

Type depositories: USNTC (holotype, paratypes), BMNH, HH, FMNH, PNM (paratypes) (Keirans and Hillyard 2001, op. cit. under H. aciculifer)

Known stages: male, female, nymph, larva

Zoogeographic Region: Oriental

Ecoregion: Luzon tropical pine forests

Hosts: usual hosts for larvae, nymphs and adult ticks are Artiodactyla: Cervidae.

Artiodactyla: Cervidae, Suidae (ANL)

Human infestation: no

Reference

Hoogstraal, H. & Parrish, D.W. 1968. Studies on Southeast Asian Haemaphysalis ticks (Ixodoidea: Ixodidae). H. (Kaiseriana.) luzonensis sp. n. from Philippine deer and boars. J. Parasitol., 54: 402–410.

90 – H. madagascariensis Colas-Belcour & Millot, 1948 (Bull. Soc. Pathol. Exot., 41: 384–388)

Type depository: IPP (holotype, paratype) (Hoogstraal, H. 1966. The Haemaphysalis ticks (Ixodoidea, Ixodidae) of birds. 2. Redescription of the type material of H. (Rhipistoma) madagascariensis Colas-Belcour and Millot. J. Parasitol., 52: 801–804), originally named H. hoodi madagascariensis.

Known stages: male, female

Zoogeographic Region: Afrotropical

Ecoregion: Madagascar subhumid forests

Hosts: Cuculiformes: Cuculidae (ANL)

Passeriformes: Vangidae (L)

Human infestation: no

Remarks: Uilenberg et al. (1979) were unable to determine the undescribed larva and nymph of this species with certainty, and we therefore consider their records provisionally valid.

Reference

Uilenberg, G., Hoogstraal, H. & Klein, J.-M. 1979. Les tiques (Ixodoidea) de Madagascar et leur rôle vecteur. Arch. Inst. Pasteur Madagascar Num. Spéc., 153 pp.

91 – H. mageshimaensis Saito & Hoogstraal, 1973 (J. Parasitol., 59: 569–578)

Type depositories: MZD (holotype, paratype), USNTC (paratypes) (Saito, Y. & Hoogstraal, H. 1973. Haemaphysalis (Kaiseriana) mageshimaensis sp. n. (Ixodoidea: Ixodidae), a Japanese deer parasite with bisexual and parthenogenetic reproduction. J. Parasitol., 59: 569–578)

Known stages: male, female, nymph, larva

Zoogeographic Regions: Oriental, Palearctic

Ecoregions: tropical and subtropical moist broadleaf forests

Hosts: Artiodactyla: Bovidae; Carnivora: Canidae, Felidae, Viverridae (ANL)

Artiodactyla: Cervidae (AN)

Artiodactyla: Suidae (A)

Rodentia: Muridae; Passeriformes: Pycnonotidae, Zosteropidae (L)

Human infestation: yes (Hoogstraal and Santana 1974)

References

Hoogstraal, H. & Santana, F.J. 1974. Haemaphysalis (Kaiseriana) mageshimaensis (Ixodoidea: Ixodidae): human and wild and domestic mammal hosts, and distribution in Japan, Taiwan, and China. J. Parasitol., 60: 866–869.

Kolonin, G.V. 1995. Review of the ixodid tick fauna (Acari: Ixodidae) of Vietnam. J. Med. Entomol., 32: 276–282.

92 – H. megalaimae Rajagopalan, 1963 (J. Parasitol., 49: 340–345)

Type depositories: USNTC (holotype, paratype), BMNH, HH, VRC, ZSI (paratypes) (Rajagopalan, P.K. 1963. Haemaphysalis megalaimae sp. n., a new tick from the small green barbet (Megalaima viridis) in India. J. Parasitol., 49: 340–345). However, Keirans and Hillyard (2001, op. cit. under H. aciculifer) do not include paratypes of this species in their list of types in the BMNH.

Known stages: male, female, nymph, larva

Zoogeographic Region: Oriental

Ecoregion: northern Indochina subtropical forests

Hosts: Piciformes: Capitonidae (ANL)

Human infestation: no

Remarks: Miranpuri and Naithani (1978) state that Haemaphysalis megalaimae has been found on birds other than Capitonidae, however without specifying the species of birds. Sun et al. (2012) determined H. megalaimae from Chrysolophus pictus (Galliformes) but we feel that this record requires confirmation. Hosts listed above are according to Rajagopalan (1972).

References

Miranpuri, G.S. & Naithani, R.C. 1978. A check-list of Indian ticks (Ixodoidea: Acarina). Indian Veterinary Research Institute, Izatnagar, 50 pp.

Rajagopalan, P.K. 1963. Haemaphysalis megalaimae sp. n., a new tick from the small green barbet (Megalaima viridis) in India. J. Parasitol., 49: 340–345.

Sun, Y., Wei, C.-C., Xu, R.-M. & Liu, X. 2012. Haemaphysalis ticks (Ixodoidea: Ixodidae) in China: subgenus Ornithophysalis, with a new record species. Acta Parasitol. Med. Entomol. Sin., 19: 50–57. In Chinese. Note: the information contained in this article has been obtained with the help of Yun, S., and we appreciate this collaboration.

93 – H. megaspinosa Saito, 1969 (Acta Med. Biol., 17: 87–96)

Type depositories: MZD (holotype, paratype), CS (paratype). (Saito, Y. 1969. Studies on ixodid ticks. X. Haemaphysalis megaspinosa n. sp. (Ixodoidea, Ixodidae) from Kanagawa Prefecture, Japan. Acta Med. Biol., 17: 87–96)

Known stages: male, female, nymph, larva

Zoogeographic Region: Palearctic

Ecoregions: temperate broadleaf and mixed forests

Hosts: usual hosts for adult ticks are Artiodactyla: Cervidae.

Artiodactyla: Cervidae; Perissodactyla: Equidae (AN)

Artiodactyla: Bovidae, Suidae; Carnivora: Canidae, Felidae, Ursidae (A)

Human infestation: yes (Sishima et al. 2000)

Remarks: Camicas et al. (1998) state that H. megaspinosa is found in the Oriental and Palearctic Regions, but we found no bona fide records for Oriental localities. The larva of H. megaspinosa is known only from laboratory-reared specimens in Yamaguti et al. (1971). Camicas et al. (1998) list pholeophilic (burrowing) mammals as hosts for the larva and nymph of H. megaspinosa, but we have found no records that support this host-parasite association. Heath (2013) lists rodents and mustelids as hosts of H. megaspinosa as a result of a transcription error (Heath, A.C.G., personal communication to Guglielmone, A.A.).

References

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. 1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.

Heath, A.C.G. 2013. Implications for New Zealand of potentially invasive ticks sympatric with Haemaphysalis longicornis Neumann, 1901 (Acari: Ixodidae). Syst. Appl. Acarol., 18: 1–26.

Kakuda, H., Shiraishi, S. & Uchida, T.A. 1989. Ticks from wild mammals in the Kyushu District including Okinawa Prefecture, Japan. J. Fac. Agr. Kyushu Univ., 33: 267–273.

Seishima, M., Izumi, T., Oyama, Z. & Kadoska, T. 2000. Tick bite by Haemaphysalis megaspinosa – First case. Eur. J. Dermatol., 10: 389–391.

Shimada, Y., Inokuma, H., Beppu, T., Okuda, M. & Onishi, T. 2003. Survey of ixodid tick species on domestic cats in Japan. Vet. Parasitol., 111: 231–239.

Takada, N., Ishiguro, F., Fujita, H., Wang, H.-P. & Masuzawa, T. 1998. Lyme disease spirochetes in ticks from northeastern China. J. Parasitol., 84: 499–504.

Yamaguti, N., Tipton, V.J., Keegan, H.L. & Toshioka, S. 1971. Ticks of Japan, Korea, and the Ryukyu Islands. Brigham Young Univ. Sci. Bull. Biol. Ser., 15 (1), 226 pp.

Yamauchi, T., Yano, S., Yamamoto, T., Yamamoto, E. & Miyamoto, T. 2013. Ticks (Acari: Ixodidae) from medium-sized to large mammals in Ehime Prefecture, Japan. Exp. Appl. Acarol., 60: 263–270.

Ybañez, A.P., Sato, F., Nambo, Y., Fukui, T., Masuzawa, T., Ohashi, N., Matsumoto, K., Kishimoto, T. & Inokuma, H. 2013. Survey on tick-borne pathogens in thoroughbred horses in the Hidaka District, Hokkaido, Japan. J. Vet. Med. Sci.,75: 11–15.

Yokohata, Y., Fujita, O., Kamiya, M., Fujita, T., Kaneko, K. & Ohbayashi, M. 1990. Parasites from the Asiatic black bear (Ursus tibetanus) on Kyushu island. J. Wildl. Dis., 26: 137–138.

94 – H. menglaensis Pang, Chen & Xiang, 1982 (Zool. Res., 3 (Suppl.): 45–51. In Chinese)

This species is not included in Camicas et al. (1998, op. cit. under H. anomaloceraea), but no reasons are given for its omission.

Type depository: IME (holotype, paratypes) (Pang, D., Chen, C. & Xiang, R. 1982. Notes on ixodid ticks from southwestern Yunnan with description of a new species of Haemaphysalis (Acarina: Ixodidae). Zool. Res., 3 (Suppl.): 45–51. In Chinese)

Known stages: male, female

Zoogeographic Region: Oriental

Ecoregion: Huang He plain mixed forests

Hosts: Artiodactyla: Cervidae (A)

Human infestation: no

References

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. 1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.

Chen, Z., Yang, X., Bu, F., Yang, X., Yang, X. & Liu, J. 2010. Ticks (Acari: Ixodoidea: Argasidae, Ixodidae) of China. Exp. Appl. Acarol., 51: 393–404.

Horak, I.G., Camicas, J.-L. & Keirans, J.E. 2002. The Argasidae, Ixodoidea and Nuttalliellidae (Acari: Ixodida): a world list of valid tick names. Exp. Appl. Acarol., 27–54.

Pang, D., Chen, C. & Xiang, R. 1982. Notes on ixodid ticks from southwestern Yunnan with description of a new species of Haemaphysalis (Acarina: Ixodidae). Zool. Res., 3 (Suppl.): 45–51. In Chinese.

95 – H. minuta Kohls, 1950 (J. Parasitol., 36: 319–321)

Type depositories: USNTC (holotype, paratypes), BMNH (paratypes) (Keirans and Hillyard 2001, op. cit. under H. aciculifer)

Known stages: male, female, nymph, larva

Zoogeographic Region: Oriental

Ecoregions: tropical and subtropical dry broadleaf forests, desert and xeric shrublands

Hosts: usual hosts for larvae, nymphs and adults are Galliformes: Phasianidae. Mammalia are considered exceptional hosts for this tick.

Galliformes: Phasianidae (ANL)

Carnivora: Viverridae (A)

Passeriformes: Timaliidae (NL)

Artiodactyla: Bovidae; Primates: Cercopithecidae; Rodentia: Hystricidae; Cuculiformes: Cuculidae; Passeriformes (several families) (N)

Rodentia: Muridae (L)

Human infestation: no

Remarks: Camicas et al. (1998) list the female as undescribed, but it had earlier been described by Santos Dias (1956). Trapido et al. (1964b) reported some variation among their female specimens but did not reject the description of Santos Dias (1956) and we therefore regard his description as conditionally valid. Kolonin (2009) limits the hosts of H. minuta to Aves and Rodentia: Hystricidae, while Hoogstraal and Kim (1985) and Camicas et al. (1998) record only Aves as hosts of this tick. We have found no evidence that might lead us to conclude that the few records of this species from Mammalia are erroneous; consequently, we regard these records as provisionally valid but believe that H. minuta is primarily a parasite of Aves. Rao et al. (1973) list “mongoose” as hosts for H. minuta but provide no further details, and we have therefore not included such hosts in our list.

References

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. 1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.

Hoogstraal, H. & Kim, K.C. 1985. Tick and mammal coevolution, with emphasis on Haemaphysalis. In K.C. Kim (editor), Coevolution of parasitic arthropods and mammals. John Wiley & Sons, New York, pp. 505–568.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Rajagopalan, P.K. 1972. Ixodid ticks (Acarina: Ixodidae) parasitizing wild birds in the Kyasanur Forest disease area of Shimoga District, Mysore State, India. J. Bombay Nat. Hist. Soc., 69: 55–78.

Rajagopalan, P.K. & Sreenivasan, M.A. 1981. Ixodid ticks on cattle and buffaloes in the Kyasanur Forest disease area of Karnataka State. Ind. J. Med. Res., 73: 880–889.

Rajagopalan, P.K., Patil, A.P. & Boshell, J. 1968. Ixodid ticks on their mammalian hosts in the Kyasanur Forest disease area of Mysore State, India, 1961–1964. Ind. J. Med. Res., 56: 510–526.

Rao, T.R., Dhanda, V., Bhat, H.R. & Kulkarni, S.M. 1973. A survey of haematophagous arthropods in western Himalayas, Sikkim and Hill Districts of West Bengal. A general account. Ind. J. Med. Res., 61: 1421–1461.

Santos Dias, J.A.T. 1956. Sobre duas entidades do género Haemaphysalis C.L. Koch, 1844. Mem. Estud. Mus. Zool. Univ. Coimbra (242), 9 pp.

Seneviratna, P. 1965. The Ixodoidea (ticks) of Ceylon. Parts II and III. Ceylon Vet. J., 13: 28–54.

Trapido, H., Goverdhan, M.K., Rajagopalan, P.K. & Rebello, M.J. 1964a. Ticks ectoparasitic on monkeys in the Kyasanur Forest disease area of Shimoga District, Mysore State, India. Am. J. Trop. Med. Hyg., 13: 763–772.

Trapido, H., Varma, M.G.R., Rajagopalan, P.K., Singh, K.R.P. & Rebello, M.J. 1964b. A guide to the identification of all stages of the Haemaphysalis ticks of South India. Bull. Entomol. Res., 55: 249–270.

96 – H. mjoebergi Warburton, 1926 (Parasitology, 18: 55–58)

Type depository: BMNH (lectotype) (Keirans and Hillyard 2001, op. cit. under H. aciculifer)

Known stages: male, female

Zoogeographic Region: Oriental

Ecoregions: tropical and subtropical moist broadleaf forests

Hosts: usual hosts for adult ticks are Artiodactyla: Cervidae.

Artiodactyla: Cervidae, Bovidae (A)

Human infestation: yes

References

Hoogstraal, H. & Wassef, H.Y. 1982. Haemaphysalis (Garnhamphysalis) mjoebergi: identity, structural variation and biosystematic implications, deer hosts, and distribution in Borneo and Sumatra (Ixodoidea: Ixodidae). J. Parasitol., 68: 138–144.

97 – H. montgomeryi Nuttall, 1912 (Parasitology, 5: 50–60)

Type depositories: BMNH (lectotype, paralectotypes), ZMB (paralectotype, see below) (Keirans & Hillyard, 2001, op. cit. under H. aciculifer). Moritz & Fischer, (1981, op. cit. under H. cinnabarina) refer to the specimen in ZMB as Paratypus, “cotype” [sic].

Known stages: male, female, nymph, larva

Zoogeographic Regions: Oriental, Palearctic

Ecoregions: montane grasslands and shrublands

Hosts: usual hosts for adult ticks are Artiodactyla: Bovidae.

Artiodactyla: Bovidae; Rodentia: Muridae (AN)

Artiodactyla: Camelidae, Cervidae; Carnivora: Canidae, Felidae; Perissodactyla: Equidae; Rodentia: Sciuridae; Passeriformes: Corvidae (A)

Carnívora: Herpestidae; Soricomorpha: Soricidae (N)

Carnivora (unknown families); Erinaceomorpha: Erinaceidae; Aves (unknown orders) (N and/or L)

Carnivora: Herpestidae; Galliformes: Phasianidae (tick stages unknown)

Human infestation: yes (Hoogstraal et al. 1966)

Remarks: Camicas et al. (1998) list this species as exclusively Oriental, but Hoogstraal and Kim (1985) and Chen et al. (2010) present data for Palearctic localities. Hoogstraal & Kim (1985) use the term “immatures” without specifying whether larvae, nymphs or both stages were present on hosts. We have found no bona fide records for hosts of larval H. montgomeryi.

References

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. 1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.

Chen, Z., Yang, X., Bu, F., Yang, X., Yang, X. & Liu, J. 2010. Ticks (Acari: Ixodoidea: Argasidae, Ixodidae) of China. Exp. Appl. Acarol., 51: 393–404.

Gupta, S.K. & Kumar, R. 1994. Ixodid ticks of camel in India and their control measures. Int. J. Anim. Sci., 9: 55–56.

Hoogstraal, H. & Kim, K.C. 1985. Tick and mammal coevolution, with emphasis on Haemaphysalis. In K.C. Kim (editor), Coevolution of parasitic arthropods and mammals. John Wiley & Sons, New York, pp. 505–568.

Hoogstraal, H., Trapido, H. & Kohls, G.M. 1966. Studies on Southeast Asian Haemaphysalis ticks (Ixodoidea, Ixodidae). Speciation in the H. (Kaiseriana) obesa group: H. semermis Neumann, H. obesa Larrousse, H. roubaudi Toumanoff, H. montgomeryi Nuttall, and H. hirsuta sp. n. J. Parasitol., 52: 169–191.

Kaul, H.N., Shetty, P.S., Ghalsasi, G.R. & Dhanda, V. 1990. Survey of ticks (Acarina: Ixodidae) for Crimean haemorrhagic fever virus activity in Jammu & Kashmir State, India. Ind. J. Med. Res. (A), 91: 5–8.

Mitchell, R.M. & Dick, J.A. 1978. Ectoparasites from Nepal birds. J. Bombay Nat. Hist. Soc., 74: 264–274.

Sharma, B.D. 1993. Medical & veterinary arthropod-disease ecology. Ashish Publishing House, New Delhi, 335 pp.

98 – H. moreli Camicas, Hoogstraal & El Kammah, 1972 (J. Parasitol., 58: 1185–1196)

Type depositories: USNTC (holotype, paratypes), ORSTOM (paratypes) (Camicas, J.-L., Hoogstraal, H. & El Kammah, K.M. 1972. Notes on African Haemaphysalis ticks. VIII. H. (Rhipistoma) moreli sp. n., a carnivore parasite of the H. (R.) leachi group (Ixodoidea: Ixodidae). J. Parasitol., 58: 1185–1196; Keirans & Clifford, 1984, op. cit. under H. bartelsi)

Known stages: male, female, nymph, larva

Zoogeographic Region: Afrotropical

Ecoregions: tropical and subtropical grasslands, savannas and shrublands

Hosts: usual hosts for adult ticks are Carnivora: Viverridae.

Carnivora: Viverridae (ANL)

Mammalia (several orders) (A)

Rodentia: Muridae (N)

Human infestation: no

Remarks: the above host list excludes adults, larvae and nymphs of H. moreli found in the nests of Primates: Galagidae by Camicas et al. (1972).

References

Camicas, J.-L., Hoogstraal, H. & El Kammah, K.M. 1972. Notes on African Haemaphysalis ticks. VIII. H. (Rhipistoma) moreli sp. n., a carnivore parasite of the H. (R.) leachi group (Ixodoidea: Ixodidae). J. Parasitol., 58: 1185–1196.

Cumming, G.S. 1998. Host preference in African ticks (Acari: Ixodida): a quantitative data set. Bull. Entomol. Res., 88: 379–406.

Morel, P.-C. 1976. Contribution à la connaissance de la distribution des tiques (Acariens, Ixodidae et Amblyommidae) en Afrique éthiopienne continentale. Annexe cartographique. Institut d’Élevage et de Médecine Vétérinaire des Pays Tropicaux, Paris, 326 pp.

99 – H. moschisuga Teng 1980 (Acta Zootax. Sin., 5: 144–149. In Chinese)

Type depository: IZAS (holotype, paratypes) (Teng, K.-F. 1980. Two new species of Haemaphysalis from China (Acarina: Ixodidae). Acta Zootax. Sin., 5: 144–149. In Chinese, NAMRU-3 translation 1777)

Known stages: male, female, nymph, larva

Zoogeographic Regions: Oriental, Palearctic

Ecoregions: deserts and xeric shrublands, montane grasslands and shrublands

Hosts: usual hosts for larvae, nymphs and adult ticks are Artiodactyla: Moschidae.

Artiodactyla: Moschidae (ANL)

Artiodactyla: Bovidae; Lagomorpha: Leporidae (A)

Galliformes: Phasianidae (NL)

Human infestation: no

Remarks: Camicas et al. (1998) list this species as exclusively Palearctic, but records from Yunnan Province by Teng and Jiang (1991) and Chen et al. (2010) indicate that H. moschisuga is also present in the Oriental Zoogeographic Region. The list of hosts above follows that of Teng (1984), but Teng and Jiang (1991) limit hosts to Artiodactyla: Bovidae and Moschidae. Kolonin (2009) retains the host profile of H. moschisuga according to Teng (1984), which we provisionally accept. Mihalca et al. (2011) regard this species as endangered.

References

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. 1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.

Chen, Z., Yang, X., Bu, F., Yang, X., Yang, X. & Liu, J. 2010. Ticks (Acari: Ixodoidea: Argasidae, Ixodidae) of China. Exp. Appl. Acarol., 51: 393–404.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Mihalca, A.D., Gherman, C.M. & Cozma, V. 2011. Coendangered hard-ticks: threatened or threatening? Parasit. Vectors, 4 (71), 7 pp.

Teng, K.-F. 1984. On the distribution, hosts and immature stages of Haemaphysalis moschisuga Teng (Acarina: Ixodidae). Acta Zootax. Sin., 9: 219–221. In Chinese, NAMRU-3 translation 1777.

Teng, K.-F. & Jiang, Z.-J. 1991. Economic insect fauna of China. Fasc. 39, Acari: Ixodidae. Science Press, Beijing, 355 pp. In Chinese.

100 – H. muhsamae Santos Dias, 1954 (Mem. Estad. Mus. Zool. Univ. Coimbra (225), 9 pp). Considerable difficulties attend the morphological identification of this species. Santos Dias, J.A.T. (1954. Mais uma nova espécie de carraça do género Haemaphysalis C.L. Koch, 1884, para a fauna de Moçambique. Mem. Estud. Mus. Zool. Univ. Coimbra (225), 9 pp.) described the male, female and nymph of H. muhsamae (as H. muhsami), but Santos Dias, J.A.T. (1955. Sobre uma pequena colecçao de carraças provenientes de Tete (Moçambique). Moçambique (81): 117–132) corrected himself concerning the female and nymph used in the original description, which were in fact H. leachi indica (a synonym of H. indica). He then redescribed the female and nymph of H. muhsamae from new material collected in Moçambique. Kolonin (2009, op. cit. under H. anomaloceraea) omits H. muhsamae from his list of ixodid ticks of the world but gives no reason for this omission. Haemaphysalis muhsamae is treated as provisionally valid here. See remarks below.

Type depository: VLM (holotype, paratype) (Santos Dias, J.A.T. 1962. Tipos entomológicos em colecçao no laboratorio central de patología veterinária de Lourenço Marques. An. Serv. Vet. Moçambique (8): 63–67) as Haemaphysalis muhsami

Known stages: male, female, (nymph?)

Zoogeographic Region: Afrotropical

Ecoregions: several Afrotropical ecoregions

Hosts: Aves are considered exceptional hosts for this tick.

Mammalia (several orders); Strigiformes: Strigidae; Passeriformes: Muscicapidae (A)

Human infestation: no

Remarks: the nymph is described in Santos Dias (1955) but not included in Camicas et al. (1998), probably because of the difficulties involved in correctly identifying this stage. Records of H. muhsamae adults from Aves are rare. Because it can be quite difficult to determine this species, we have not included the few records of sub-adult ticks. Walker (1991) considers the records of Theiler (1962) of H. leachii [sic] mühsami [sic] as in fact referring to H. spinulosa, and Theiler’s records have therefore not been included in the above host list. Keirans and Durden (2001) record the introduction of H. muhsamae into the Nearctic Region, but there is no evidence that it has become established there. With the exception of Keirans and Durden (2001), all authors listed below refer to this species as Haemaphysalis leachi muhsami.

References

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. 1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.

Elbl, A. & Anastos, G. 1966. Ixodid ticks (Acarina, Ixodidae) of Central Africa. Vol. IV. Genera Aponomma Neumann, 1899, Boophilus Curtice, 1891, Dermacentor Koch, 1844, Haemaphysalis Koch, 1844, Hyalomma Koch, 1844 and Rhipicentor Nuttall and Warburton, 1908. Ann. Mus. R. Centr. Afr., Ser. 8º Sci. Zool. (148), 412 pp.

Hoogstraal, H. 1956. African Ixodoidea. I. Ticks of the Sudan (with special reference to Equatoria Province and with preliminary reviews of the genera Boophilus, Margaropus and Hyalomma). Research Report NM 005 050.29.07, Department of the Navy Bureau of Medicine and Surgery, Washington D.C. 1101 pp.

Keirans, J.E. & Durden, L.A. 2001. Invasion: exotic ticks (Acari: Argasidae, Ixodidae) imported into the United States. A review and new records. J. Med. Entomol., 38: 850–861.

Santos Dias, J.A.T. 1955. Sobre uma pequena colecçao de carraças provenientes de Tete (Moçambique). Moçambique (81): 117–132

Theiler, G. 1962. The Ixodoidea parasites of vertebrates in Africa south of the Sahara (Ethiopian Region). Report to the Director of Veterinary Services, Onderstepoort, South Africa, Project S.9958, 260 pp.

Walker, J.B. 1991. A review of the ixodid ticks (Acari, Ixodidae) occurring in southern Africa. Onderstepoort J. Vet. Res., 58: 81–105.

101 – H. nadchatrami Hoogstraal, Trapido & Kohls, 1965 (J. Parasitol., 51: 433–451)

Type depositories: USNTC (holotype, paratypes), BMNH, FMNH, HH, IMRKL, MCZ, MNHN, UM, ZMB (paratypes) (Keirans and Hillyard 2001, op. cit. under H. aciculifer) as H. papuana nadchatrami.

Known stages: male, female, nymph

Zoogeographic Region: Oriental

Ecoregions: tropical and subtropical moist broadleaf forests

Hosts: Rodentia: Muridae (ANL)

Artiodactyla: Bovidae, Suidae; Carnivora: Canidae, Felidae, Mustelidae; Perissodactyla: Equidae, Tapiridae (A)

Artiodactyla: Tragulidae (NL)

Carnivora: Viverridae; Rodentia: Sciuridae (N)

Human infestation: yes (Tanskul et al. 1983)

Remarks: hosts for the undescribed larva of H. nadchatrami are recorded in Hoogstraal et al. (1972), and we regard these records as provisionally valid. Tanskul et al. (1983) identified adults of H. nadchatrami from Muridae. This host family is ignored for imagoes of H. nadchatrami in Kolonin (2009) but we consider this record provisionally valid.

References

Hoogstraal, H., Trapido, H. & Kohls, G.M. 1965. Southeast Asian Haemaphysalis ticks (Ixodoidea, Ixodidae). H. (Kaiseriana) papuana nadchatrami ssp. n. and redescription of H. (K.) semermis Neumann. J. Parasitol., 51: 433–451.

Hoogstraal, H., Lim, B.L., Nadchatram, M. & Anastos, G. 1972. The Gunong Benom Expedition 1967. 8. Ticks (Ixodidae) of Gunong Benom and their altitudinal distribution, hosts and medical relationships. Bull. Br. Mus. (Nat. Hist.) Zool., 23: 167–186.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Tanskul, P., Stark, H.E. & Inlao, I. 1983. A checklist of ticks of Thailand (Acari: Metastigmata: Ixodoidea). J. Med. Entomol., 20: 330–341.

102 – H. nepalensis Hoogstraal, 1962 (J. Parasitol., 48: 195–203)

Type depositories: USNTC (holotype, paratype), HH (paratype) (Hoogstraal, H. 1962. Haemaphysalis nepalensis sp. n. from a Himalayan rodent and man, and description of the male of H. aponommoides Warburton (n. comb.) (Ixodoidea, Ixodidae). J. Parasitol., 48: 195–203)

Known stages: male, female, nymph

Zoogeographic Regions: Oriental, Palearctic

Ecoregions: montane grasslands and shrublands

Hosts: Artiodactyla: Bovidae (AN)

Carnivora: Felidae, Ursidae; Rodentia (unknown family) (A)

Human infestation: yes (Hoogstraal 1962)

Remarks: Camicas et al. (1998) list this species as exclusively Oriental, but there are also bona fide records for the Palearctic Region. Mitchell (1979) lists Carnivora: Ailuridae and Soricomorpha: Soricidae as hosts of H. nepalensis without stating which stages of this tick were found on these hosts, and we have therefore tentatively excluded them from our host list. Hoogstraal and Kim (1985) and Kolonin (2009) state that the immature stages of H. nepalensis feed on the same hosts as adults, but apart from nymphs found on Bovidae by Dhanda (1964) we were unable to confirm this. Mitchell and Dick (1978) present a record of H. nepalensis from Aves (tick stages unknown), but Aves are not included in Hoogstraal and Kim (1985) and are not included in our list above.

References

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. 1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.

Chen, Z., Yang, X., Bu, F., Yang, X., Yang, X. & Liu, J. 2010. Ticks (Acari: Ixodoidea: Argasidae, Ixodidae) of China. Exp. Appl. Acarol., 51: 393–404.

Dhanda, V. 1964. Haemaphysalis nepalensis Hoogstraal, 1962 (Ixodoidea: Ixodidae), systematic position based on description of the nymph, and new host and locality records. J. Parasitol., 50: 783–785.

Hoogstraal, H. 1962. Haemaphysalis nepalensis sp. n. from a Himalayan rodent and man, and description of the male of H. aponommoides Warburton (n. comb.) (Ixodoidea, Ixodidae). J. Parasitol., 48: 195–203.

Hoogstraal, H. & Kim, K.C. 1985. Tick and mammal coevolution, with emphasis on Haemaphysalis. In K.C. Kim (editor), Coevolution of parasitic arthropods and mammals. John Wiley & Sons, New York, pp. 505–568.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Mitchell, R.M. 1979. A list of ectoparasites from Nepalese mammals, collected during the Nepal ectoparasite program. J. Med. Entomol., 16: 227–233.

Mitchell, R.M. & Dick, J.A. 1978. Ectoparasites from Nepal birds. J. Bombay Nat. Hist. Soc., 74: 264–274.

Teng, K.-F. & Jiang, Z.-J. 1991. Economic insect fauna of China. Fasc. 39, Acari: Ixodidae. Science Press, Beijing, 355 pp. In Chinese.

103 – H. nesomys Hoogstraal, Uilenberg & Klein, 1966 (J. Parasitol., 52: 1199–1202)

Type depository: USNTC (holotype) (Keirans & Clifford, 1984, op. cit. under H. bartelsi)

Known stages: male

Zoogeographic Region: Afrotropical

Ecoregion: Madagascar subhumid forests

Hosts: Rodentia: Nesomyidae (A)

Human infestation: no

Reference

Uilenberg, G., Hoogstraal, H. & Klein, J.-M. 1979. Les tiques (Ixodoidea) de Madagascar et leur rôle vecteur. Arch. Inst. Pasteur Madagascar Num. Spéc., 153 pp.

104 – H. norvali Hoogstraal & Wassef, 1983 (Onderstepoort J. Vet. Res., 50: 183–189)

Type depositories: USNTC (holotype, paratypes), VRLH, OVI, ORSTOM (paratypes) (Hoogstraal, H. & Wassef, H.Y. 1983. Notes on African Haemaphysalis ticks. XV. H.

(Rhipistoma) norvali sp. n., a hedgehog parasite of the H. (R.) spinulosa group in Zimbabwe (Acarina: Ixodidae). Onderstepoort J. Vet. Res., 50: 183–189)

Known stages: male, female, nymph, larva

Zoogeographic Region: Afrotropical

Ecoregion: Southern Africa bushveld

Hosts: Erinaceomorpha: Erinaceidae (A)

Human infestation: no

Remarks: the larva and nymph of H. norvali are known only from laboratory-reared specimens (Hoogstraal and Wassef 1983).

Reference

Hoogstraal, H. & Wassef, H.Y. 1983. Notes on African Haemaphysalis ticks. XV. H. (Rhipistoma) norvali sp. n., a hedgehog parasite of the H. (R.) spinulosa group in Zimbabwe (Acarina: Ixodidae). Onderstepoort J. Vet. Res., 50: 183–189.

105 – H. novaeguineae Hirst, 1914 (Trans. R. Soc. Lond., 20: 325–334)

This name should not be confused with Haemaphysalis novaguineae Krijgsman and Ponto, 1932, a synonym of H. bancrofti as explained in Roberts (1963, op. cit. under H. bremneri).

Type depository: BMNH (lectotype, paralectotypes) (Keirans and Hillyard 2001, op. cit. under H. aciculifer)

Known stages: male, female, nymph, larva

Zoogeographic Region: Australasian

Ecoregions: tropical and subtropical moist broadleaf forests; tropical and subtropical grasslands, savannas and shrublands

Hosts: Artiodactyla: Cervidae, Suidae; Peramelemorphia: Peramelidae (ANL)

Diprotodontia: Macropodidae (AN)

Cuculiformes: Cuculidae (AL)

Carnivora: Canidae; Monotremata: Tachyglossidae; Perissodactyla: Equidae; Rodentia: Muridae; Gruiformes: Rallidae (A)

Carnivora: Felidae (N)

Artiodactyla: Bovidae; Chiroptera: Pteropodidae (stage unknown)

Human infestation: yes (Unsworth et al. 2007)

Remarks: Camicas et al. (1998) state that H. novaeguineae is found in the Australasian and Oriental Zoogeographic Regions, but we follow Hoogstraal and Kim (1985), who categorized H. novaeguineae as an exclusively Australasian species. Although Hoogstraal (1982) found several specimens of H. novaeguineae on Aves, these records are ignored by Kolonin (2009).

References

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. 1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.

Hoogstraal, H. 1982. Ticks (Acari: Ixodoidea): a preliminary study. Monogr. Biol., 42: 537–544.

Hoogstraal, H. & Kim, K.C. 1985. Tick and mammal coevolution, with emphasis on Haemaphysalis. In K.C. Kim (editor), Coevolution of parasitic arthropods and mammals. John Wiley & Sons, New York, pp. 505–568.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Owen, I.L. 2011. Parasites of animals in Papua New Guinea recorded at the National Veterinary Laboratory: a catalogue, historical review and zoogeographical affiliations. Zootaxa, 3143: 1–163.

Roberts, F.H.S. 1963. A systematic study of the Australian species of the genus Haemaphysalis Koch (Acarina: Ixodidae). Aust. J. Zool., 11: 35–80.

Roberts, F.H.S. 1969, The larvae of Australian Ixodidae (Acarina: Ixodoidea). J. Aust. Entomol. Soc., 8: 37–78.

Roberts, F.H.S. 1970. Australian ticks. CSIRO, Melbourne, 267 pp.

Unsworth, N.B., Stenos, J., Graves, S.R., Faa, A.G., Cox, G.E., Dyer, J.R., Boutlis, C.S., Lane, A.M., Shaw, M.D., Robson, J. & Nisse, M.D. 2007. Flinders Island spotted fever rickettsioses caused by a “marmionii” strain of Rickettsia honei, eastern Australia. Emerg. Inf. Dis., 13: 566–572.

106 – H. obesa Larrousse, 1925 (Ann. Parasitol. Hum. Comp., 3: 301–305)

Type depository: IP (lectotype) (Hoogstraal et al. 1966, op. cit. under H. hirsuta)

Known stages: male, female, nymph, larva

Zoogeographic Region: Oriental

Ecoregions: tropical and subtropical dry broadleaf forests

Hosts: Mammalia (several orders) (A)

Carnivora: Canidae, Mustelidae (N and/or L)

Human infestation: yes (Tanskul et al. 1983)

Remarks: Tanskul et al. (1983) and Hoogstraal and Kim (1985) use the term “immatures” without indicating whether larvae, nymphs or both stages were found on hosts; consequently, the hosts for the larvae and nymphs of H. obesa are not known with certainty.

References

Hoogstraal, H. & Kim, K.C. 1985. Tick and mammal coevolution, with emphasis on Haemaphysalis. In K.C. Kim (editor), Coevolution of parasitic arthropods and mammals. John Wiley & Sons, New York, pp. 505–568.

Hoogstraal, H., Saito, Y., Dhanda, V. & Bhat, H.R. 1971. Haemaphysalis (H.) obesa Larrousse (Ixodoidea: Ixodidae) from Northeast India and Southeast Asia: description of immature stages and biological observations. J. Parasitol., 57: 177–184.

Tanskul, P., Stark, H.E. & Inlao, I. 1983. A checklist of ticks of Thailand (Acari: Metastigmata: Ixodoidea). J. Med. Entomol., 20: 330–341.

107 – H. obtusa Dönitz, 1910 (Denkschr. Med.-Naturw. Ges. Jena, 16: 397–494)

See remarks below.

Type depositories: ZMB, BMNH (syntypes) (Moritz and Fischer 1981, op. cit. under H. cinnabarina; Keirans and Hillyard 2001, op. cit. under H. aciculifer)

Known stages: male, female, nymph

Zoogeographic Region: Afrotropical

Ecoregion: Madagascar subhumid forests

Hosts: usual hosts for adult ticks are Carnivora: Eupleridae and Viverridae.

Carnivora: Eupleridae (ANL)

Carnivora: Viverridae (AN)

Canidae (A)

Afrosoricida: Tenrecidae; Rodentia: Muridae, Nesomyidae (NL)

Human infestation: no

Remarks: the larva of H. obtusa has not been described. Uilenberg et al. (1979) feel that determinations of nymphs and larvae from Afrosoricida and Muridae, in the absence of adult ticks, should be considered tentative. We too regard these records as only provisionally valid. Furthermore, Uilenberg et al. (1979) state that there are “typical” and “atypical” specimens of H. obtusa, thus implying that more than one species may be represented under this name.

Reference

Uilenberg, G., Hoogstraal, H. & Klein, J.-M. 1979. Les tiques (Ixodoidea) de Madagascar et leur rôle vecteur. Arch. Inst. Pasteur Madagascar Num. Spéc., 153 pp.

108 – H. oliveri Apanaskevich & Horak, 2008 (J. Parasitol., 94: 594–607)

Some specimens of H. oliveri collected before its description were identified as members of the H. leachi group, as noted in Apanaskevich and Horak (2008, op. cit. under H. colesbergensis). Kolonin (2009, op. cit. under H. anomaloceraea) does not include this species in his list of Ixodidae of the world.

Type depositories: USNTC (holotype, paratypes), OVI, ZIAC (paratypes) (Apanaskevich and Horak 2008, op. cit. under H. colesbergensis)

Known stages: male, female, larva

Zoogeographic Region: Afrotropical

Ecoregions: Sahelian acacia savanna; east Sudanian savanna

Hosts: usual hosts for adult ticks are Carnivora: Canidae and Felidae.

Carnivora: Canidae, Felidae, Viverridae; Artiodactyla: Bovidae (A)

Human infestation: no

Remarks: the larva of H. oliveri is known only from laboratory-reared specimens (Apanaskevich and Horak 2008).

Reference

Apanaskevich, D.A. & Horak, I.G. 2008. Two new species of African Haemaphysalis ticks (Acari: Ixodidae), carnivore parasites of the H. (Rhipistoma) leachi group. J. Parasitol., 94: 594–607.

109 – H. orientalis Nuttall & Warburton, 1915 (Ticks. A monograph of the Ixodoidea. Part III. The genus Haemaphysalis. Cambridge University Press, London, pp. 349–550)

See remarks below.

Type depository: BMNH (lectotype, paralectotypes) (Keirans and Hillyard 2001, op. cit. under H. aciculifer), originally named H. hoodi orientalis.

Known stages: male, female

Zoogeographic Region: Afrotropical

Ecoregions: tropical and subtropical grasslands, savannas and shrublands

Hosts: usual hosts for adult ticks are Hyracoidea: Procaviidae.

Artiodactyla: Bovidae; Hyracoidea: Procaviidae (A)

Human infestation: no

Remarks: Santos Dias (1953) refers to this tick as Haemaphysalis zambeziae Santos Dias, 1953, which Hoogstraal (1956) classifies as a synonym of H. orientalis. Although Santos Dias later (1962) recognized this synonymy, he continued to maintain his earlier (1953) assertion that Bovidae are hosts of this species, which Hoogstraal (1956) believes to be an error, as inferred from correspondence between Santos Dias and Theiler, (quoted on page 168 of Hoogstraal 1956). Therefore, our acceptance of this host-parasite relationship in the above list is provisional. Kolonin (2009) excludes Bovidae as hosts for H. orientalis.

References

Hoogstraal, H. 1956. Notes on African Haemaphysalis ticks. III. The hyrax parasites, H. bequaerti sp. nov., H. orientalis N. and W., 1915 (new combination), and H. cooleyi Bedford, 1929 (Ixodoidea, Ixodidae). J. Parasitol., 42: 156–172.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Santos Dias, J.A.T. 1953. Sobre uma nova espécie de carraça do gênero Haemaphysalis Koch, 1844, (Acarina; Ixodoidea) para a fauna de Moçambique. Mem. Estud. Mus. Zool. Univ. Coimbra (219), 7 pp.

Santos Dias, J.A.T. 1962. Tipos entomológicos em colecçao no laboratorio central de patología veterinária de Lourenço Marques. An. Serv. Vet. Moçambique (8): 63–67.

110 – H. ornithophila Hoogstraal & Kohls, 1959 (J. Parasitol., 45: 417–420)

Type depositories: MCZ (holotype, paratypes), HH, USNTC (paratypes) (Hoogstraal, H. & Kohls, G.M. 1959. The Haemaphysalis ticks (Acarina, Ixodidae) of birds. I. H. ornithophila n. sp. from Burma and Thailand. J. Parasitol., 45: 417–420)

Known stages: male, female

Zoogeographic Regions: Oriental, Palearctic

Ecoregions: northern triangle subtropical forests; lower Gangetic plains moist deciduous forests

Hosts: Galliformes: Phasianidae (ANL)

Passeriformes: Turdidae (AN)

Passeriformes: Pittidae, Chloropseidae; Carnivora: Mustelidae; Lagomorpha: Leporidae; Artiodactyla: Bovidae (A)

Human infestation: no

Remarks: we have been unable to find descriptions of the larva and nymph of H. ornithophila. Wilson (1970), however, discusses several records of these stages from Aves that we consider tentatively valid. Kolonin (1995) states that specimens of H. ornithophila found on Chloropseidae are equivalent to H. bacthaiensis Phan Trong, 1977. Mitchell (1979) regards Cervidae as hosts of H. ornithophila, but this view has not been corroborated in subsequent works (Kim et al. 2009).

References

Chen, Z., Yang, X., Bu, F., Yang, X., Yang, X. & Liu, J. 2010. Ticks (Acari: Ixodoidea: Argasidae, Ixodidae) of China. Exp. Appl. Acarol., 51: 393–404.

Hoogstraal, H. & Kohls, G.M. 1959. The Haemaphysalis ticks (Acarina, Ixodidae) of birds. I. H. ornithophila n. sp. from Burma and Thailand. J. Parasitol., 45: 417–420.

Hoogstraal, H. & Wassef, H.Y. 1973. The Haemaphysalis ticks (Ixodoidea: Ixodidae) of birds. 3.H. (Ornithophysalis) subgen. n.: definition, species, hosts and distribution in the Oriental, Palearctic, Malagasy, and Ethiopian Faunal Regions. J. Parasitol., 59: 1099–1117.

Kang, J.-G., Kim, H.-C., Choi, C.-Y., Nam, H.-Y., Chae, H.-Y., Chong, S.-T., Klein, T.A., Ko, S. & Chae, J.-S. 2013. Molecular detection of Anaplasma, Bartonella, and Borrelia species in ticks collected from migratory birds from Hong-do Island, Republic of Korea. Vector- Borne Zoon. Dis., 13: 215–225.

Kim, H.C., Ko, S., Choi, C.-Y., Nam, H.-Y., Chae, H.Y., Chong, S.T., Klein, T.A., Sames, W.J., Robbins, R.G. & Chae, J.-S. 2009. Migratory bird tick surveillance, including a new record of Haemaphysalis ornithophila Hoogstraal and Kohls 1959 (Acari: Ixodidae) from Hong-do (Hong Island), Republic of Korea. Syst. Appl. Acarol., 14: 3–10.

Kolonin, G.V. 1995. Review of the ixodid tick fauna (Acari: Ixodidae) of Vietnam. J. Med. Entomol., 32: 276–282.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Mitchell, R.M. 1979. A list of ectoparasites from Nepalese mammals, collected during the Nepal ectoparasite program. J. Med. Entomol., 16: 227–233.

Teng, K.-F. & Jiang, Z.-J. 1991. Economic insect fauna of China. Fasc. 39, Acari: Ixodidae. Science Press, Beijing, 355 pp. In Chinese.

Wilson, N. 1970. New distributional records of ticks from Southeast Asia and the Pacific (Metastigmata: Argasidae, Ixodidae). Oriental Insects, 4: 37–46.

Xu, R. & Li, K. 1997. A collection of ticks from Guizhou, China. Syst. Appl. Acarol., 2: 245–246.

111 – H. palawanensis Kohls, 1950 (Natl. Inst. Health Bull. (192), 28 pp)

Type depositories: CNHM (holotype, paratypes), USNTC, PNM (paratypes) (Kohls 1950, op. cit. under H. hoogstraali)

Known stages: male, female

Zoogeographic Region: Oriental

Ecoregion: Palawan rain forests

Hosts: Carnivora (several families); Rodentia: Hystricidae, Muridae, Sciuridae; Artiodactyla: Suidae, Cervidae (A)

Human infestation: no

Remarks: Kohls (1950) found a female H. palawanensis crawling on a human, but we have not included humans in our host list above. Cervids are not listed as hosts of this tick in Kolonin (2009), but there is a valid record for this type of host in Kohls (1950). Durden and Keirans (1996) regard H. palawanensis as an endangered species.

References

Durden, L.A. & Keirans, J.E. 1996. Host-parasite coextinction and the plight of tick conservation. Am. Entomol., 42: 87–91.

Kohls, G.M. 1950. Ticks (Ixodoidea) of the Philippines. Natl. Inst. Health Bull. (192), 28 pp.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

112 – H. papuana Thorell, 1883 (Ann. Mus. Civ. Stor. Nat. Genova, 18: 21–69)

Type depository: GM (syntypes) (Anastos 1950, op. cit. under H. hylobatis)

Known stages: male, female, nymph, larva

Zoogeographic Regions: Australasian, Oriental

Ecoregions: tropical and subtropical moist broadleaf forests

Hosts: usual hosts for adult ticks are Artiodactyla: Suidae. Aves are considered exceptional hosts for this tick.

Artiodactyla: Suidae; Carnivora: Canidae (AN)

Carnivora: Ursidae (AL)

Rodentia: Muridae (A, N and/or L)

Artiodactyla: Cervidae, Moschidae; Carnivora: Felidae, Mustelidae, Viverridae; Rodentia: Hystricidae; Galliformes: Phasianidae (A)

Passeriformes: Muscicapidae (stage unknown)

Human infestation: yes (Tanskul et al. 1983)

Remarks: Camicas et al. (1998) list the larva of H. papuana as undescribed, but it was earlier described in Kadarsan (1971). Reports of this species published prior to Trapido et al. (1964), who refer to it as H. papuana papuana, have not been included in our analyses because of diagnostic uncertainties. Tanskul et al. (1983) use the term “immatures” without specifying whether they found larvae, nymphs or both stages on hosts. Yamaguti et al. (1971) state that reports of this tick from Korea, which lies within the Palearctic Region, are erroneous. According to Keirans (1985), a larva of H. papuana from Ursidae should be considered tentative, and we have therefore provisionally included it in our host list above. Kolonin (2009) excludes Muridae as hosts for this tick, but we accept records from these hosts in Tanskul et al. (1983) and also an odd record from Aves in Gould et al. (1970).

References

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. 1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.

Durden, L.A., Merker, S. & Beati, L. 2008. The tick fauna of Sulawesi, Indonesia (Acari: Ixodoidea: Argasidae and Ixodidae). Exp. App. Acarol., 45: 85–110.

Gould, D.J., Clifford, C.M., Lakshana, P., Emerson, K.C., Hoogstraal, H., Marshall. J.T., Stark, H.E., Nadchatram, M., Davis, E.W., Inlao, I., Maneechai, N., Klaimanee, N. & Punthusiri, P. 1970. Distribution and ecology of ectoparasites of vertebrates in Southeast Asia. Annual Program Report SEATO Medical Research Laboratory, April 1969-March 1970, United States Army Medical Component, Bangkok, Thailand, pp, 97–105.

Hoogstraal, H. 1982. Ticks (Acari: Ixodoidea): a preliminary study. Monogr. Biol., 42: 537–544.

Kadarsan, S. 1971. Larval ixodid ticks of Indonesia (Acarina: Ixodidae). Ph.D. Dissertation, University of Maryland, 182 pp.

Keirans, J.E. 1985. George Henry Falkiner Nuttall and the Nuttall tick catalogue. U. S. Dept. Agric., Agric. Res. Ser. Misc. Pub. (1438), 1785 pp.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Phan Trong, C. 1977. Ve bet va con trung ky sinh o Viet Nam. Tap 1. Ve (Ixodoidea), mo ta va phan loai. Ha Noi: Khoa hoc va ky thuat, 489 pp. In Vietnamese.

Tanskul, P., Stark, H.E. & Inlao, I. 1983. A checklist of ticks of Thailand (Acari: Metastigmata: Ixodoidea). J. Med. Entomol., 20: 330–341.

Trapido, H., Hoogstraal, H. & Varma, M.G.R. 1964. Status and descriptions of Haemaphysalis p. papuana Thorell (n. comb.) and of H. papuana kinneari Warburton (n. comb.) (Ixodoidea, Ixodidae) of southern Asia and New Guinea. J. Parasitol., 50: 172–188.

Yamaguti, N., Tipton, V.J., Keegan, H.L. & Toshioka, S. 1971. Ticks of Japan, Korea, and the Ryukyu Islands. Brigham Young Univ. Sci. Bull. Biol. Ser., 15 (1), 226 pp.

113 – H. paraleachi Camicas, Hoogstraal & El Kammah, 1983 (J. Parasitol., 69: 400–404)

See remarks below

Type depositories: USNTC (holotype, paratypes), ORSTOM, CM (paratypes) (Camicas, J.-L., Hoogstraal, H. & El Kammah, K.M. 1983. Notes on African Haemaphysalis ticks. XIV. Description of adults of H. (Rhipistoma) paraleachi sp. n., a carnivore parasite of the H. (R.) leachi group (Ixodoidea: Ixodidae). J. Parasitol., 69: 400–404)

Known stages: male, female

Zoogeographic Region: Afrotropical

Ecoregions: several Afrotropical ecoregions

Hosts: usual hosts for adult ticks are Carnivora: Canidae, Felidae and Viverridae.

Rodentia: Muridae (AN)

Carnivora (several families); Artiodactyla: Bovidae; Primates: Cercopithecidae (A)

Human infestation: yes (El Kammah et al. 1992)

Remarks: Kolonin (2009) does not list hosts for nymphs of this tick, probably because the nymph has not been described, but we provisionally accept the records of this stage in Cornet (1995). Prior to its description, H. paraleachi was confused with H. leachi by several authors (Camicas et al. 1983).

References

Camicas, J.-L., Hoogstraal, H. & El Kammah, K.M. 1983. Notes on African Haemaphysalis ticks. XIV. Description of adults of H. (Rhipistoma) paraleachi sp. n., a carnivore parasite of the H. (R.) leachi group (Ixodoidea: Ixodidae). J. Parasitol., 69: 400–404

Cornet, J.-P. 1995. Contribution à l’étude des tiques (Acarina: Ixodina) de la République Centrafricaine 4. Inventaire et répartition. Acarologia, 36: 203–212.

El Kammah, K.M., Hoogstraal, H. & Camicas, J.-L. 1992. Notes on African ticks: XI. H. (Rhipistoma) paraleachi (Ixodoidea: Ixodidae) distribution and hosts of adults. Int. J. Acarol., 18: 205–212.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

114 – H. paraturturis Hoogstraal, Trapido & Rebello, 1963 (J. Parasitol., 49: 686–691)

Type depositories: USNTC (holotype, paratypes), BMNH, VRC, IM (paratypes) (Keirans and Hillyard 2001, op. cit. under H. aciculifer). This species was originally identified as H. bispinosa intermedia Warburton & Nuttall, 1909, a synonym of H. intermedia, by Nuttall & Warburton (1915, op. cit. under H. celebensis), as noted in Hoogstraal. H., Trapido, H. & Rebello, M.J. (1963. Haemaphysalis paraturturis sp. n., a carnivore parasite of the H. turturis group in India (Ixodoidea, Ixodidae). J. Parasitol., 49: 686–691).

Known stages: male, female, nymph

Zoogeographic Region: Oriental

Ecoregion: lower Gangetic Plains moist deciduous forests

Hosts: Carnivora: Canidae, Felidae, Ursidae; Artiodactyla: Bovidae, Suidae (A)

Cuculiformes: Cuculidae (N)

Rodentia: Muridae, Sciuridae; Soricomorpha: Soricidae; Passeriformes: Timaliidae (N and/or L)

Human infestation: no

Remarks: Kolonin (2009) does not list Rodentia, Soricomorpha and Passeriformes as hosts for this tick, but we provisionally accept records from these hosts reported by Geevarghese and Dhanda (1995), who do not specify whether their collections contained larvae, nymphs or both stages.

References

Geevarghese, G. & Dhanda, V. 1995. Ixodid ticks of Maharashtra State, India. Acarologia, 36: 309–313.

Hoogstraal. H., Trapido, H. & Rebello, M.J. 1963. Haemaphysalis paraturturis sp. n., a carnivore parasite of the H. turturis group in India (Ixodoidea, Ixodidae). J. Parasitol., 49: 686–691.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

115 – H. parmata Neumann, 1905 (Arch. Parasitol., 9: 225–241)

Type depository: BMNH (syntypes) (Keirans and Hillyard 2001, op. cit. under H. aciculifer)

Known stages: male, female, nymph, larva

Zoogeographic Region: Afrotropical

Ecoregions: tropical and subtropical moist broadleaf forests; few ticks in tropical and subtropical grasslands, savannas and shrublands; montane grassland and shrublands

Hosts: usual hosts for adult ticks are Artiodactyla: Bovidae. We consider Squamata exceptional hosts for this tick, while Aves are considered exceptional hosts for adult ticks.

Mammalia (several orders); Galliformes: Phasianidae, Numididae (ANL)

Squamata: Elapidae (A)

Passeriformes: Strildidae (N)

Passeriformes: Viduidae; Squamata: Atractaspididae (L)

Ciconiiformes: Bucorvidae (N and/or L)

Human infestation: yes (Cornet 1995)

Remarks: Theiler (1962) uses the term “immatures” without specifying whether larvae, nymphs or both preimaginal stages of H. parmata were found on hosts. Infestations of adult ticks on Aves and any stage of development on Squamata appear to be infrequent events and have been ignored in Kolonin (2009). We, however, consider the occasional records from these hosts in Keirans (1985) and Morel (2003), among others, provisionally valid.

References

Aeschlimann, A. 1967. Biologie et écologie des tiques (Ixodoidea) de Côte d’Ivoire. Acta Trop., 24: 281–405.

Cornet, J.-P. 1995. Contribution à l’étude des tiques (Acarina: Ixodina) de la République Centrafricaine 4. Inventaire et répartition. Acarologia, 36: 203–212.

Elbl, A. & Anastos, G. 1966. Ixodid ticks (Acarina, Ixodidae) of Central Africa. Vol. IV. Genera Aponomma Neumann, 1899, Boophilus Curtice, 1891, Dermacentor Koch, 1844, Haemaphysalis Koch, 1844, Hyalomma Koch, 1844 and Rhipicentor Nuttall and Warburton, 1908. Ann. Mus. R. Centr. Afr., Ser. 8º Sci. Zool. (148), 412 pp.

Horak, I.G., Boomker, J. & Flamand, J.R.B. 1991. Ixodid ticks and lice infesting red duikers and bushpigs in north-eastern Natal. Onderstepoort J. Vet. Res., 58: 281–284.

Keirans, J.E. 1985. George Henry Falkiner Nuttall and the Nuttall tick catalogue. U. S. Dept. Agric., Agric. Res. Ser. Misc. Pub. (1438), 1785 pp.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Morel, P.-C. 2003. Les tiques d’Afrique et du Bassin méditerranéen (1965–1995). CIRAD- EMVT, 1342 pp.

Theiler, G. 1962. The Ixodoidea parasites of vertebrates in Africa south of the Sahara (Ethiopian Region). Report to the Director of Veterinary Services, Onderstepoort, South Africa, Project S.9958, 260 pp.

Walker, J.B. 1974. The ixodid ticks of Kenya. A review of present knowledge of their hosts and distribution. Commonwealth Institute of Entomology, London, 220 pp.

Walker, J.B. 1991. A review of the ixodid ticks (Acari, Ixodidae) occurring in southern Africa. Onderstepoort J. Vet. Res., 58: 81–105.

116 – H. parva (Neumann, 1897) (Mém. Soc. Zool. Fr., 10: 324–420)

This species was originally designated as Dermacentor parvus, but Morel, P.-C. (1963. Sur quelques Haemaphysalis paléarctiques (Acariens, Ixodoidea). Ann. Parasitol. Hum. Comp., 38: 915–924) recognized that it was in fact a member of the genus Haemaphysalis and identical to H. otophila Schulze, 1919. Therefore, D. parvus and H. otophila schulzei Tonelli-Rondelli, 1926 are synonyms of H. parva. Neumann, L.G. (1908. Notes sur les ixodidés. VI. Arch. Parasitol., 12: 5–27) described H. parva from Ceylon (Sri Lanka); however, the name was preoccupied and the correct name for the species described by Neumann (1908) is H. intermedia Warburton & Nuttall, 1909, as stated in Trapido, H. & Hoogstraal, H. (1963. Status of Haemaphysalis bispinosa var. intermedia Warburton & Nuttall, 1909, next available name for H. parva Neumann, 1908 (preoccupied) (Ixodoidea, Ixodidae). J. Parasitol., 49: 691–692). See also remarks below.

Type depository: MNHN (holotype) (Neumann, 1897, op. cit. under H. flava) as Dermacentor parvus

Known stages: male, female, nymph, larva

Zoogeographic Region: Palearctic

Ecoregions: mediterranean forests, woodlands and scrub

Hosts: Testudines are considered exceptional hosts for this tick, while Aves are considered exceptional hosts for adult ticks.

Rodentia: Sciuridae; Lagomorpha: Leporidae (AN)

Erinaceomorpha: Erinaceidae (AL)

Artiodactyla: Bovidae; Carnivora: Mustelidae; Falconiformes: Falconidae; Passeriformes: Sturnidae (A, N and/or L)

Several orders (Mammalia) (A)

Galliformes: Phasianidae (AL)

Rodentia: Cricetidae, Spalacidae; Charadriiformes: Scolopacidae; Squamata: Lacertidae (NL)

Rodentia: Dipodidae, Muridae; Falconiformes: Accipitridae; Gruiformes: Gruidae; Squamata: Colubridae (N)

Passeriformes: Muscicapidae (L)

Charadriiformes: Burhinidae, Charadriidae; Passeriformes (several families); Piciformes: Picidae (N and/or L)

Lagomorpha: Ochotonidae; Rodentia: Gliridae, Calomyscidae; Squamata: Viperidae; Testudines: Testudinidae (stages unknown)

Human infestation: yes (Bursali et al. 2012)

Remarks: Camicas et al. (1998) state that the larva of H. parva is undescribed, but there is a description in Filippova (1997). Ter-Vartanov et al. (1954), Hoogstraal (1959), Feider (1964), Theodor and Costa (1967) and Yeruham et al. (1996) refer to this species as H. otophila, while Inci et al. (2003) use the names of H. otophila and H. parva simultaneously for ticks parasitizing Bovidae. Hoogstraal & Kim (1985) use the term “immatures” without specifying larvae, nymphs or both, while Morel (2003) presents information on larvae and nymphs of H. parva, but it is difficult to infer their hosts from his data. Faizi et al. (2011) recorded infestations of Squamata: Scincidae with adults, nymphs and larvae of H. parva, but we believe that these records need confirmation and we have not included them in our host list above. Kolonin (2009) records only Mammalia as hosts for this tick.

References

Aksn, N. & Aksn, N.E. 2002. The prevalence of ectoparasites on wild rabbits in Elazg Region, Turkey. Turk. Parazitol. Derg., 26: 67–70. In Turkish.

Bursali, A., Keskin, A. & Tekin, S. 2012. A review of the ticks (Acari: Ixodida) of Turkey: species diversity, hosts and geographical distribution. Exp. Appl. Acarol., 57: 91–104.

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. 1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.

Faizi, H., Rastegar-Pouyani, N. & Yarani, R. 2011. On the occurrence of ectoparasite ticks on Trachylepis and Eumeces (Reptilia: Scincidae) in Iran. Amphib. Reptile Conserv., 5: 7–10.

Fedorov, V.G. 1972. Ticks of the superfamily Ixodoidea of birds and their nests in western Siberia. In A. Cherepanov & L.A. Mater (editors), Transcontinental associations of migratory birds and their role in distribution of arboviruses. 5. Simp. Isuch. Roli Pereletn. Ptits Rasp. Arbovirus, Novosibirsk, Jul. 1969, pp. 384–388. In Russian, NAMRU-3 translation 1494.

Feider, Z. 1964. Les connaissances actuelles sur les Acariens de Roumanie (Ixodides, Gamasides et Trombiculides). Acarologia (Fasc. Hors Sér.): 262–274.

Filippova, N.A. 1997. Ixodid ticks of subfamily Amblyomminae. Fauna of Russia and neighbouring countries, 4 (5), Nauka, St. Petersburg, 436 pp. In Russian.

Hoogstraal, H. 1959. Biological observations on certain Turkish Haemaphysalis ticks (Ixodoidea, Ixodidae). J. Parasitol., 45: 227–232.

Hoogstraal, H. & Kim, K.C. 1985. Tick and mammal coevolution, with emphasis on Haemaphysalis. In K.C. Kim (editor), Coevolution of parasitic arthropods and mammals. John Wiley & Sons, New York, pp. 505–568.

Inci, A., Nalbantoğlu, S., Çam, Y., Atasever, A., Karaer, Z., Çakmak, A., Sayin, F., Yukari, B.A., Iça, A. & Deniz, A. 2003. Theileriosis and tick infestations in sheep and goats around Kayseri. Turk J. Vet. Anim. Sci., 27: 57–60. In Turkish.

Keysary, A., Eremeeva, M.E., Leitner, M., Din, A.B., Wikswo, M.E., Mumcuoglu, K.Y., Inbar, M., Wallach, A.D., Shanas, U., King, R. & Waner, T. 2011. Spotted fever group rickettsiae collected from wild animals in Israel. Am. J. Trop. Med. Hyg., 85: 919–923.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Morel, P.-C. 2003. Les tiques d’Afrique et du Bassin méditerranéen (1965–1995). CIRAD- EMVT, 1342 pp.

Ter-Vartanov, V.H., Gusev, V.H., Bakeev, N.N., Labunets, N.F., Guseva, A.A. & Reznik, P.A. 1954. On the question of transmission of mammalian ectoparasites by birds. Zool. Zh., 33: 1116–1125. In Russian, NAMRU-3 translation 52.

Theodor, O. & Costa, M. 1967. A survey of the parasites of wild mammals and birds in Israel. Part one. Ectoparasites. The Israel Academy of Science and Humanities, Jerusalem, 119 pp.

Yeruham, I., Hadani, A., Galker, F. & Rosen, S. 1996. The seasonal occurrence of ticks (Acari: Ixodidae) on sheep and in the field in the Judean area of Israel. Exp. Appl. Acarol., 20: 47–56.

117 – H. pavlovskyi Pospelova-Shtrom, 1935 (Trudy Tadzhik. Bazy Akad. Nauk SSSR (5): 205–217. In Russian)

Camicas et al. (1998, op. cit. under H. anomaloceraea) and Kolonin (2009, op. cit. under H. anomaloceraea) consider this species a synonym of H. doenitzi. Guglielmone et al. (2009, op. cit. under H. colasbelcouri) discussed the status of this species, concluding that it is valid, and we concur. See also H. phasiana.

Type depository: ZIAC (holotype, paratypes) (Filippova 2008, op. cit. under H. caucasica)

Known stages: male, female, nymph, larva

Zoogeographic Region: Palearctic

Ecoregions: desert and xeric shrublands

Hosts: Galliformes: Phasianidae; Lagomorpha: Leporidae (ANL)

Passeriformes: Sylviidae (L)

Human infestation: no

Remarks: see H. doenitzi.

References

Filippova, N.A. 1997. Ixodid ticks of subfamily Amblyomminae. Fauna of Russia and neighbouring countries, 4 (5), Nauka, St. Petersburg, 436 pp. In Russian.

Hoogstraal, H., Kaiser, M.N., Traylor, M.A., Guindy, E. & Gaber, S. 1963. Ticks (Ixodidae) on birds migrating from Europe and Asia to Africa, 1959–61. Bull. World Health. Org., 28: 236–262.

118 – H. pedetes Hoogstraal, 1972 (J. Parasitol., 58: 979–983)

Type depository: BMNH (holotype, paratype) (Keirans and Hillyard 2001, op. cit. under H. aciculifer), originally identified as H. cooleyi. See also H. cooleyi.

Known stages: male, female

Zoogeographic Region: Afrotropical

Ecoregions: tropical and subtropical grasslands, savannas and shrublands

Hosts: usual hosts for adult ticks are Rodentia: Pedetidae.

Rodentia: Pedetidae (ANL)

Carnivora: Felidae, Mustelidae, Viverridae (A)

Human infestation: no

Remarks: we were unable to find descriptions of the larva and nymph of H. pedetes, but we tentatively accept records of nymphs and larvae on Pedetidae (Anderson and Kok 2003), although these have not been included in Kolonin (2009). Cumming (1998) lists Rodentia: Pedetidae as the only hosts for this tick, but Hoogstraal and Kim (1985) record parasitism of Carnivora. See also H. cooleyi.

References

Anderson, P.C. & Kok, O.B. 2003. Ectoparasites of springhares in the Northern Cape Province, South Africa. S. Afr. J. Wildl. Res., 33: 23–32.

Cumming, G.S. 1998. Host preference in African ticks (Acari: Ixodida): a quantitative data set. Bull. Entomol. Res., 88: 379–406.

Hoogstraal, H. & Kim, K.C. 1985. Tick and mammal coevolution, with emphasis on Haemaphysalis. In K.C. Kim (editor), Coevolution of parasitic arthropods and mammals. John Wiley & Sons, New York, pp. 505–568.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Walker, J.B. 1991. A review of the ixodid ticks (Acari, Ixodidae) occurring in southern Africa. Onderstepoort J. Vet. Res., 58: 81–105.

119 – H. pentalagi Pospelova-Shtrom, 1935 (Trudy Tadzhik. Bazy Akad. Nauk SSSR (5): 205–217. In Russian)

Type depository: ZIAC (holotype) (Filippova 2008, op. cit. under H. caucasica)

Known stages: male, female, nymph, larva

Zoogeographic Region: Oriental

Ecoregion: Nansei Islands subtropical evergreen forests

Hosts: Lagomorpha: Leporidae (ANL)

Human infestation: no

Remarks: Camicas et al. (1998) state that this tick is found in the Oriental and Palearctic Zoogeographic Regions, but we found no bona fide records of H. pentalagi from the Palearctic. Mihalca et al. (2011) regard this species as endangered.

References

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. 1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.

Hoogstraal, H. & Kim, K.C. 1985. Tick and mammal coevolution, with emphasis on Haemaphysalis. In K.C. Kim (editor), Coevolution of parasitic arthropods and mammals. John Wiley & Sons, New York, pp. 505–568.

Kitaoka, S. & Suzuki, H. 1974. Reports of medico-zoology investigations in the Nansei Islands. Part 2. Ticks and their seasonal prevalences in southern Amami-oshima. Jap. J. Sanit. Zool., 26: 21–26. In Japanese.

Mihalca, A.D., Gherman, C.M. & Cozma, V. 2011. Coendangered hard-ticks: threatened or threatening? Parasit. Vectors, 4 (71), 7 pp.

120 – H. petrogalis Roberts, 1970 (Australian ticks. CSIRO, Melbourne, 267 pp)

Type depositories: QM (holotype, paratypes), ANIC, AM, USNTC (paratypes) (Roberts, F.H.S. 1970. Australian ticks. CSIRO, Melbourne, 267 pp.; Halliday, B. personal communication to Guglielmone, A.A.)

Known stages: male, female

Zoogeographic Region: Australasian

Ecoregion: Queensland tropical rain forests

Hosts: Diprotodontia: Macropodidae (A)

Human infestation: no

Reference

Roberts, F.H.S. 1970. Australian ticks. CSIRO, Melbourne, 267 pp.

121 – H. phasiana Saito, Hoogstraal & Wassef, 1974 (J. Parasitol., 60: 198–208)

Kolonin (2009, op. cit. under H. anomaloceraea) treats H. phasiana, as well as H. pavlovskyi, as synonyms of H. doenitzi. This decision was based on morphological differences ascribed to colonization of the periphery of the geographic range of H. doenitzi. We follow Guglielmone et al. (2010, op. cit. under H. anomaloceraea), maintaining that more convincing evidence is needed to conclude that H. phasiana and H. pavlovskyi are synonyms of H. doenitzi. See also H. doenitzi and remarks below.

Type depositories: USNTC (holotype, paratypes), CS (paratypes) (Saito, Y., Hoogstraal, H. & Wassef, H.Y. 1974. The Haemaphysalis ticks (Ixodoidea: Ixodidae) of birds. 4. H. (Ornithophysalis) phasiana sp. n. from Japan. J. Parasitol., 60: 198–208)

Known stages: male, female, nymph, larva

Zoogeographic Regions: Oriental, Palearctic

Ecoregions: temperate broadleaf and mixed forests

Hosts: Galliformes: Phasianidae (ANL)

Passeriformes: Emberizidae; Artiodactyla: Cervidae (AN)

Passeriformes: Muscicapidae; Lagomorpha: Leporidae (A)

Gruiformes: Turnicidae; Passeriformes: Sylviidae (N)

Passeriformes: Emberizidae (stage unknown)

Human infestation: no

Remarks: Sames et al. (2008) believe that more than one taxon is represented under the name H. phasiana. Hoogstraal and Kim (1985) state that H. phasiana is an exclusive parasite of Aves; however, there are about 20 records for this species (Sames et al. 2008), and three of them refer to ticks found on mammals. We therefore conclude that more information is needed to determine whether H. phasiana is indeed an exclusive parasite of Aves and whether records from Mammalia are exceptional. See also H. doenitzi.

References

Chen, Z., Yang, X., Bu, F., Yang, X., Yang, X. & Liu, J. 2010. Ticks (Acari: Ixodoidea: Argasidae, Ixodidae) of China. Exp. Appl. Acarol., 51: 393–404.

Filippova, N.A. 1997. Ixodid ticks of subfamily Amblyomminae. Fauna of Russia and neighbouring countries, 4 (5), Nauka, St. Petersburg, 436 pp. In Russian.

Hoogstraal, H. & Kim, K.C. 1985. Tick and mammal coevolution, with emphasis on Haemaphysalis. In K.C. Kim (editor), Coevolution of parasitic arthropods and mammals. John Wiley & Sons, New York, pp. 505–568.

Sames, W.J., Kim, H.C., Chong, S.T., Lee, I.Y., Apanaskevich, D.A., Robbins, R.G., Bast. J., Moore, R. & Klein, T.A. 2008. Haemaphysalis (Ornithophysalis) phasiana (Acari: Ixodidae) in the Republic of Korea: two province records and habitat descriptions. Syst. Appl. Acarol., 13: 43–50.

Yamauchi, T. 2001. A bibliographical survey of host-parasite relationships between birds and ticks from Japan. Bull. Hoshizaki Green Found., 5: 271–308. In Japanese.

122 – H. pospelovashtromae Hoogstraal, 1966 (J. Parasitol., 52: 787–800)

See H. danieli.

Type depositories: USNTC (holotype, paratypes), ZMB (paratype) (Moritz and Fischer 1981, op. cit. under H. cinnabarina; Keirans and Clifford 1984, op. cit. under H. bartelsi). One paratype was previously determined as H. warburtoni by Hoogstraal, H. (1966. Haemaphysalis (Allophysalis) pospelovashtromae sp. n. from USSR and redescription of the type material of H. (A.) warburtoni Nuttall from China (Ixodoidea, Ixodidae). J. Parasitol., 52: 787–800). See “remarks on some invalid names” for a new synonym of H. pospelovashtromae.

Known stages: male, female, nymph, larva

Zoogeographic Region: Palearctic

Ecoregions: desert and xeric shrublands

Hosts: Artiodactyla: Bovidae, Suidae; Rodentia: Sciuridae (A)

Mammalia (several orders); Passeriformes: Sittidae (NL)

Human infestation: no

Remarks: see H. warburtoni.

Reference

Filippova, N.A. 1997. Ixodid ticks of subfamily Amblyomminae. Fauna of Russia and neighbouring countries, 4 (5), Nauka, St. Petersburg, 436 pp. In Russian.

123 – H. primitiva Teng, 1982 (Acta Zootax. Sin., 7: 46–48. In Chinese)

Type depository: IZAS probable (holotype) (Teng, K.-F. 1982. On the subgenus Alloceraea of genus Haemaphysalis from China, with description of a new species (Ixodoidea: Ixodidae). Acta Zootax. Sin., 7: 46–48. In Chinese, translation RR1 by Robbins, F.-M.Y.; Keirans and Robbins 1999, op. cit. under H. demidovae)

Known stages: male, female

Zoogeographic Region: Oriental

Ecoregions: temperate broadleaf and mixed forests

Hosts: unknown

Human infestation: no

References

Keirans, J.E. & Robbins, R.G. 1999. A world checklist of genera, subgenera, and species of ticks (Acari: Ixodida) published from 1973 to 1997. J. Vector Ecol., 24: 115–129.

Teng, K.-F. 1982. On the subgenus Alloceraea of genus Haemaphysalis from China, with description of a new species (Ixodoidea: Ixodidae). Acta Zootax. Sin., 7: 46–48. In Chinese, translation RR1 by Robbins, F.-M.Y.

124 – H. psalistos Hoogstral, Kohls & Parrish, 1967 (J. Parasitol., 53: 1096–1102)

Type depositories: USNTC (holotype, paratypes), BMNH, BM, FMNH, HH (paratypes) (Keirans & Hillyard, 2001, op. cit. under H. aciculifer)

Known stages: male, female, nymph

Zoogeographic Regions: Australasian, Oriental

Ecoregions: Luzon tropical pine forests; Sulawesi montane rain forests

Hosts: Artiodactyla: Cervidae (AN)

Artiodactyla: Bovidae, Suidae (A)

Rodentia: Muridae (NL)

Human infestation: no

Remarks: Camicas et al. (1998) and Kolonin (2009) state that this species is exclusively Oriental, but the records of Durden et al. (2008) for Sulawesi Island belong to the Australasian Region. Kolonin (2009) does not consider Bovidae hosts for H. psalistos, but bona fide records of this relationship are reported in Durden et al. (2008). The latter authors treat their record of the undescribed larva of this tick on Muridae as tentative, and we have therefore provisionally included murids in our host list for H. psalistos. Mihalca et al. (2011) regard this species as endangered.

References

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. 1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.

Durden, L.A., Merker, S. & Beati, L. 2008. The tick fauna of Sulawesi, Indonesia (Acari: Ixodoidea: Argasidae and Ixodidae). Exp. App. Acarol., 45: 85–110.

Hoogstraal, H., Kohls, G.M. & Parrish, D.W. 1967. Studies on Southeast Asian Haemaphysalis ticks (Ixodoidea, Ixodidae). H. (Kaiseriana) psalistos sp. n., a parasite of deer and boars in Luzon, Philippines. J. Parasitol., 53: 1096–1102.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Mihalca, A.D., Gherman, C.M. & Cozma, V. 2011. Coendangered hard-ticks: threatened or threatening? Parasit. Vectors, 4 (71), 7 pp.

125 – H. punctaleachi Camicas, Hoogstraal & El Kammah, 1973 (J. Parasitol., 59: 563–568)

Guglielmone et al. (2010, op. cit. under H. anomalocerae) erroneously cite the year of description as 1983.

Type depositories: USNTC (holotype, paratype), ORSTOM (paratypes) (Camicas, J.-L., Hoogstraal, H. & El Kammah, K.M. 1973. Notes on African Haemaphysalis ticks. XI. H. (Rhipistoma) punctaleachi sp. n., a parasite of West African forest carnivores (Ixodoidea: Ixodidae). J. Parasitol., 59: 563–568)

Known stages: male, female

Zoogeographic Region: Afrotropical

Ecoregions: central Congolian lowland forests; Guinean lowland forests

Hosts: usual hosts for adult ticks are Carnivora: Viverridae.

Artiodactyla: Bovidae; Carnivora (several families); Rodentia: Hystricidae (A)

Human infestation: no

Remarks: Kolonin (2009) ignores Bovidae as hosts of H. punctaleachi, but there is a valid record for this type of host in Camicas et al. (1973).

References

Camicas, J.-L., Hoogstraal, H. & El Kammah, K.M. 1973. Notes on African Haemaphysalis ticks. XI. H. (Rhipistoma) punctaleachi sp. n., a parasite of West African forest carnivores (Ixodoidea: Ixodidae). J. Parasitol., 59: 563–568.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Pourrut, X., Emane, K.A., Camicas, J.-L., Leroy, E. & González, J.-P. 2011. Contribution to the knowledge of ticks (Acarina: Ixodidae) in Gabon. Acarologia, 51: 465–471.

126 – H. punctata Canestrini & Fanzago, 1878 (Atti R. Ist. Veneto Sci Lett. Arti (1877–1878), Ser. 5, 4: 69–208)

Type depositories: the types are probably lost (Nuttall and Warburton 1915, op. cit. under H. celebensis). These authors designate this species as Haemaphysalis cinnabarina punctata.

Known stages: male, female, nymph, larva

Zoogeographic Region: Palearctic

Ecoregions: several Palearctic ecoregions

Hosts: Testudines and Squamata: Viperidae are considered exceptional hosts for this tick.

Mammalia (several orders); Aves (several orders) (ANL)

Squamata: Anguidae, Lacertidae (NL)

Squamata: Viperidae; Testudines: Testudinidae (stages unknown)

Human infestation: yes (Burridge 2011)

Remarks: there are records of introductions of H. punctata into the Nearctic Region but no evidence that it has become established there (Burridge 2011). Ekanem et al. (2012) reported a natural population of H. punctata in the Afrotropical Region, but this is regarded as an error in identification. Kolonin (2009) ignores hosts other than mammals and birds, but, although rare, there are several valid records of H. punctata on Squamata and Testudines in the references listed below. See also H. cinnabarina.

References

Burridge, M.J. 2011. Non-native and invasive ticks. Threats to human and animal health in the United States. University of Florida Press, Gainesville, 448 pp.

Chen, Z., Yang, X., Bu, F., Yang, X., Yang, X. & Liu, J. 2010. Ticks (Acari: Ixodoidea: Argasidae, Ixodidae) of China. Exp. Appl. Acarol., 51: 393–404.

Ekanem, M.S., Opara, K.N., Bennie, D.P. & Udo, I.O. 2012. Ectoparasites [sic] of goats in Etinan, Akwa Ibom State, Nigeria. J. Agric. Biol. Sci., 2: 108–113.

Filippova, N.A. 1997. Ixodid ticks of subfamily Amblyomminae. Fauna of Russia and neighbouring countries, 4 (5), Nauka, St. Petersburg, 436 pp. In Russian.

Garben, A.F.M., Vos, H. & Von Bronswijk, J.E.M.H. 1981. Haemaphysalis punctata Canestrini & Fanzago 1877 [sic], a tick of pastured seadunes on the island of Texel (The Netherlands). Acarologia, 23: 19–25.

Grebenyuk, R.V. 1966. Ixodid ticks (Parasitiformes, Ixodidae) of Kirgizia. Akademii Nauk Kirgizkoy SSR, Instituta Biologii, Frunze, 328 pp. In Russian.

Keirans, J.E. & Durden, L.A. 2001. Invasion: exotic ticks (Acari: Argasidae, Ixodidae) imported into the United States. A review and new records. J. Med. Entomol., 38: 850–861.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Neumann, L.G. 1901. Révision de la famille des ixodidés (4ª mémorie). Mém. Soc. Zool. Fr., 14: 249–372.

Nosek, J. 1971. The ecology, bionomics, and behaviour of Haemaphysalis (Aboimisalis) punctata tick in central Europe. Z. Parasitenkd., 37: 198–210.

Sartbayev, S.K. 1959. Materials on the biology of certain ticks of the genus Haemaphysalis in Kirgizia SSR. Trud. Inst. Zool. Parasitol. Akad. Nauk Kirgiz SSR (7): 191–202. In Russian, NAMRU-3 translation 201.

Ter-Vartanov, V.H., Gusev, V.H., Bakeev, N.N., Labunets, N.F., Guseva, A.A. & Reznik, P.A. 1954. On the question of transmission of mammalian ectoparasites by birds. Zool. Zh., 33: 1116–1125. In Russian, NAMRU-3 translation 52.

127 – H. quadriaculeata Kolonin, 1992 (In V.E. Sokolov (editor), Zoological Researches in Vietnam. Nauka, Moscow, pp. 242–277. In Russian)

Type depository: ZIAC (holotype, paratype) (Filippova 2008, op. cit. under H. caucasica)

Known stages: male, female

Zoogeographic Region: Oriental

Ecoregion: south China-Vietnam subtropical evergreen forests

Hosts: Carnivora: Canidae, Mustelidae; Rodentia: Spalacidae (A)

Human infestation: no

Remarks: Kolonin (2003) states that the supposed illustration of the male of H. darjeeling in Tanskul and Inlao (1989) is closer to H. quadriaculeata than to H. darjeeling. See also H. darjeeling.

References

Kolonin, G.V. 2003. New data on ixodid tick fauna of Vietnam. Entomol. Rev., 83 (Suppl. 2): S190-S192.

Tanskul, P. & Inlao, I. 1989. Keys to the adult ticks of Haemaphysalis Koch, 1844, in Thailand with notes on changes in taxonomy (Acari: Ixodoidea: Ixodidae). J. Med. Entomol., 26: 573–601.

128 – H. quinghaiensis Teng, 1980 (Acta Zootax. Sin., 5: 144–149. In Chinese)

This name is written as H. qinghaiensis in the original description, but Camicas et al. (1998, op. cit. under H. anomaloceraea) note that the correct Latin spelling is H. quinghaiensis and we agree.

Type depository: IZAS (Keirans and Robbins 1999, op. cit. under H. demidovae) as Haemaphysalis qinghaiensis.

Known stages: male, female, nymph, larva

Zoogeographic Regions: Oriental, Palearctic

Ecoregions: temperate broadleaf and mixed forests

Hosts: Artiodactyla: Bovidae; Perissodactyla: Equidae; Lagomorpha: Leporidae (ANL)

Human infestation: yes

Remarks: Camicas et al. (1998) state that H. quinghaiensis is found exclusively in the Oriental Region; however, most records in Teng and Jiang (1991) are from Palearctic localities. Camicas et al. (1998) regard ungulates as the only significant hosts for this tick species, but Teng and Cui (1984) stress the importance of Leporidae as hosts for H. quinghaiensis.

References

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. 1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.

Chen, Z., Yang, X., Bu, F., Yang, X., Yang, X. & Liu, J. 2010. Ticks (Acari: Ixodoidea: Argasidae, Ixodidae) of China. Exp. Appl. Acarol., 51: 393–404.

Teng, K.-F. & Cui, Y.-Q. 1984. Biological observations and descriptions of immature stages of Haemaphysalis qinghaiensis Teng. Acta Entomol. Sin., 27: 330–333. In Chinese, NAMRU-3 translation 1778.

Teng, K.-F. & Jiang, Z.-J. 1991. Economic insect fauna of China. Fasc. 39, Acari: Ixodidae. Science Press, Beijing, 355 pp. In Chinese.

129 – H. ramachandrai Dhanda, Hoogstraal & Bhat, 1970 (J. Parasitol., 56: 823–831)

Type depositories: VRC (holotype, paratypes), BMNH, IM, HH, USNTC, ZIAC (paratypes) (Keirans and Hillyard 2001, op. cit. under H. aciculifer; Filippova 2008, op. cit. under H. caucasica).

Known stages: male, female, nymph, larva

Zoogeographic Region: Oriental

Ecoregion: Upper Gangetic plains moist deciduous forests

Hosts: Artiodactyla: Cervidae; Carnivora: Felidae (ANL)

Artiodactyla: Bovidae (A)

Human infestation: yes (Dhanda et al. 1970)

Reference

Dhanda, V., Hoogstraal, H. & Bhat, H.R. 1970. Haemaphysalis (Kaiseriana) ramachandrai sp.n. (Ixodoidea, Ixodidae), a parasite of man and domestic and wild mammals in northern India and Nepal. J. Parasitol., 56: 823–831.

130 – H. ratti Kohls, 1948 (J. Parasitol., 34: 154–157)

Type depositories: USNTC (holotype, paratypes), AM, BMNH (paratypes) (Keirans and Hillyard 2001, op. cit. under H. aciculifer)

Known stages: male, female, nymph, larva

Zoogeographic Region: Australasian

Ecoregions: tropical and subtropical grasslands, savannas and shrublands

Hosts: Rodentia: Muridae (ANL)

Dasyuromorphia: Dasyuridae (AL)

Peramelemorphia: Peramelidae; Diprotodontia: Phalangeridae (A)

Human infestation: no

Remarks: Kolonin (2009) states that “small marsupials” are hosts for all stages of H. ratti, but we have been unable to confirm this statement.

References

Bennett, M.D., Woolford, L., Banazis, M.J., O’Hara, J., Warren, K.S., Nicholls, P.K., Sims, C. & Fenwick, S.G. 2011. Coxiella burnetti in western barred bandicoots (Perameles bougainville) from Bernier and Dorre Islands in Western Australia. Ecohealth, 8: 519–524.

Hoogstraal, H. 1982. Ticks (Acari: Ixodoidea): a preliminary study. Monogr. Biol., 42: 537–544.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Lorch, D., Fisher, D.O. & Spratt, D.M. 2007. Variation in ectoparasite infestation on the brown antechinus Antechinus stuartii, with regard to host, habitat and environmental parameters. Aust. J. Zool., 55: 169–176.

Roberts, F.H.S. 1963. A systematic study of the Australian species of the genus Haemaphysalis Koch (Acarina: Ixodidae). Aust. J. Zool., 11: 35–80.

Roberts, F.H.S. 1970. Australian ticks. CSIRO, Melbourne, 267 pp.

131 – H. renschi Schulze, 1933 (Arch. Hydrobiol. Suppl., 12, 4: 490–502)

Type depository: USNTC (neotype) (Keirans & Clifford, 1984, op. cit. under H. bartelsi)

Known stages: male, female, nymph, larva

Zoogeographic Regions: Australasian, Oriental

Ecoregions: Sumatra and Java rain forest; Lesser Sundas deciduous forests

Hosts: Artiodactyla: Bovidae, Cervidae; Perissodactyla: Equidae (ANL)

Artiodactyla: Suidae; Carnivora: Canidae (A)

Charadriiformes: Sternidae (N)

Rodentia: Muridae (L)

Human infestation: no

Remarks: Camicas et al. (1998) maintain that H. renschi is a strictly Oriental species, but the records of Durden et al. (2008) for Sulawesi Island indicate that this tick is also present in the Australasian Region. Additionally, Durden et al. (2008) recorded the larva of H. renschi from Muridae, but this, as well as the record from Aves in Hoogstraal and Anastos (1968), is ignored in Kolonin (2009).

References

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. 1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.

Durden, L.A., Merker, S. & Beati, L. 2008. The tick fauna of Sulawesi, Indonesia (Acari: Ixodoidea: Argasidae and Ixodidae). Exp. Appl. Acarol., 45: 85–110.

Hoogstraal, H. & Anastos, G. 1968. Studies on Southeast Asian Haemaphysalis ticks (Ixodoidea: Ixodidae). Redescription of H. (Kaiseriana) renschi Schulze (resurrected), and its hosts and distribution in Indonesia. J. Parasitol., 54: 1214–1222.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

132 – H. roubaudi Toumanoff, 1940 (Rev. Méd. Fr. Extr.-Orient, 18: 463–490)

Type depositories: IPP, USNTC (syntypes) (Hoogstraal et al. 1966, op. cit. under H. hirsuta)

Known stages: male

Zoogeographic Region: Oriental

Ecoregion: southeastern Indochina dry evergreen forests

Hosts: Artiodactyla: Cervidae (A)

Human infestation: yes (Kolonin 1995)

References

Hoogstraal, H., Trapido, H. & Kohls, G.M. 1966. Studies on Southeast Asian Haemaphysalis ticks (Ixodoidea, Ixodidae). Speciation in the H. (Kaiseriana) obesa group: H. semermis Neumann, H. obesa Larrousse, H. roubaudi Toumanoff, H. montgomeryi Nuttall, and H. hirsuta sp. n. J. Parasitol., 52: 169–191.

Kolonin, G.V. 1995. Review of the ixodid tick fauna (Acari: Ixodidae) of Vietnam. J. Med. Entomol., 32: 276–282.

133 – H. rugosa Santos Dias, 1956 (Mem. Estud. Mus. Zool. Univ. Coimbra (242), 9 pp.)

Type depository: BMNH (holotype) (Hoogstraal, H. & El Kammah, K.M. 1972. Notes on African Haemaphysalis ticks. X. H. (Kaiseriana) aciculifer Warburton and H. (K.) rugosa Santos Dias, the African representatives of the spinigera subgroup (Ixodoidea: Ixodidae). J. Parasitol., 58: 960–978; Keirans, J.E. 1985. George Henry Falkiner Nuttall and the Nuttall tick catalogue. U. S. Dept. Agric., Agric. Res. Ser. Misc. Pub. (1438), 1785 pp.), as H. aciculifer rugosa, originally identified as H. aciculifer, as stated in Santos Dias, J.A.T. (1956. Notas ixodológicas. Sobre duas entidades do género Haemaphysalis C.L. Koch, 1844. Mem. Estud. Mus. Zool. Univ. Coimbra (242), 9 pp.). Surprisingly, Keirans and Hillyard (2001, op. cit. under H. aciculifer) do not record the presence of the holotype of H. rugosa in BMNH.

Known stages: male, female, nymph, larva

Zoogeographic Region: Afrotropical

Ecoregions: tropical and subtropical grasslands, savannas and shrublands

Hosts: usual hosts for adult ticks are Artiodactyla: Bovidae.

Carnivora: Viverridae (ANL)

Carnivora: Herpestidae (AN)

Artiodactyla: Bovidae (A)

Lagomorpha: Leporidae; Rodentia: Muridae (N)

Human infestation: no

Remarks: we regard records by Camicas (1978) of H. rugosa from Muridae and Leporidae as valid, but Cumming (1998) seemingly does not recognize these. A record of H. rugosa from Felidae that was considered questionable by Hoogstraal and El-Kammah (1972) has not been included in the host list above. Cornet (1995) makes a general statement, which we have been unable to confirm, that ungulates and carnivores are specific hosts for all parasitic stages of H. rugosa, but he is probably confusing H. aciculifer with H. rugosa. Cornet (1995), supported by Rousselot (1951), states that Canidae are hosts of H. rugosa. However, the latter author lists Canidae as hosts of H. aciculifer, not H. rugosa. We have been unable to find records of larvae and nymphs of H. rugosa from any ungulate host and have excluded them from the list above.

References

Camicas, J.-L. 1978. Contribution à l’étude des tiques du Sénégal (Acarida: Ixodida) 2. Description des stades préimaginales d’Haemaphysalis (Kaiseriana) rugosa Santos Dias, 1956. Cah. ORSTOM Sér. Entomol. Méd. Parasitol., 16: 23–28.

Cornet, J.-P. 1995. Contribution à l’étude des tiques (Acarina: Ixodina) de la République Centrafricaine 4. Inventaire et répartition. Acarologia, 36: 203–212.

Cumming, G.S. 1998. Host preference in African ticks (Acari: Ixodida): a quantitative data set. Bull. Entomol. Res., 88: 379–406.

Hoogstraal, H. & El Kammah, K.M. 1972. Notes on African Haemaphysalis ticks. X. H. (Kaiseriana) aciculifer Warburton and H. (K.) rugosa Santos Dias, the African representatives of the spinigera subgroup (Ixodoidea: Ixodidae). J. Parasitol., 58: 960–978.

Rousselot, R. 1951. Ixodes de l’Afrique noire. Bull. Soc. Pathol. Exot., 44: 307–309.

Walker, J.B. 1991. A review of the ixodid ticks (Acari, Ixodidae) occurring in southern Africa. Onderstepoort J. Vet. Res., 58: 81–105.

134 – H. rusae Kohls, 1950 (Natl. Inst. Health Bull. (192), 28 pp.)

Type depository: CNHM (holotype, paratype), USNTC (holotype) (Kohls, 1950, op. cit. under H. hoogstraali)

Known stages: male, female, nymph

Zoogeographic Regions: Australasian, Oriental

Ecoregions: tropical and subtropical moist broadleaf forests

Hosts: usual hosts for adult ticks are Artiodactyla: Cervidae and Suidae.

Artiodactyla: Suidae (AN)

Artiodactyla: Cervidae; Diprotodontia: Macropodidae (A)

Human infestation: no

Remarks: Hoogstraal and Wassef (1983) state that the presence of this tick species in the Australasian Region is a result of its hosts having entered this region from the Oriental Region. This is highly speculative and we provisionally consider the Australasian Region to be part of the natural range of H. rusae.

Reference

Hoogstraal, H. & Wassef, H.Y. 1983. Haemaphysalis (Garnhamphysalis) rusae (Ixodoidea: Ixodidae): identity, deer and pig hosts, and distribution in Luzon and Mindanao. J. Parasitol., 69: 215–220.

135 – H. sambar Hoogstraal, 1971 (J. Parasitol., 57: 173–176)

Type depository: BMNH (holotype) (Keirans and Hillyard 2001, op. cit. under H. aciculifer). The holotype of this species is in the Nuttall Collection, held in the BMNH, and was previously identified as H. campanulata (Hoogstraal, H. 1971. Haemaphysalis (H.) sambar sp. n. (Ixodoidea: Ixodidae), a parasite of the sambar deer in southern India. J. Parasitol., 57: 173–196).

Known stages: male

Zoogeographic Region: Oriental

Ecoregion: south western Ghats montane rain forests

Hosts: Artiodactyla: Cervidae (A)

Human infestation: no

Remarks: Mihalca et al. (2011) regard H. sambar as an endangered species.

References

Hoogstraal, H. 1971. Haemaphysalis (H.) sambar sp. n. (Ixodoidea: Ixodidae), a parasite of the sambar deer in southern India. J. Parasitol., 57: 173–196.

Mihalca, A.D., Gherman, C.M. & Cozma, V. 2011. Coendangered hard-ticks: threatened or threatening? Parasit. Vectors, 4 (71), 7 pp.

136 – H. sciuri Kohls, 1950 (Natl. Inst. Health Bull. (192), 28 pp.)

Type depositories: CNHM (holotype, paratype), USNTC (paratype) (Kohls 1950, op. cit. under H. hoogstraali)

Known stages: male

Zoogeographic Region: Oriental

Ecoregion: Mindanao-eastern visayas rain forests

Hosts: Rodentia: Sciuridae (A)

Human infestation: no

Reference

Kohls, G.M. 1950. Ticks (Ixodoidea) of the Philippines. Natl. Inst. Health Bull. (192), 28 pp.

137 – H. semermis Neumann, 1901 (Mém. Soc. Zool. Fr., 14: 249–372)

Type depository: MNHN (holotype) (Trapido, 1965, op. cit. under H. lagrangei)

Known stages: male, female, nymph

Zoogeographic Region: Oriental

Ecoregions: Sumatra and Java rain forests; Lesser Sundas deciduous forests

Hosts: Artiodactyla: Tragulidae; Rodentia: Muridae (ANL)

Carnivora: Canidae; Viverridae (AN)

Artiodactyla: Cervidae, Suidae; Carnivora: Felidae, Ursidae; Perissodactyla: Tapiridae (A)

Rodentia: Sciuridae; Scandentia: Tupaiidae (N)

Human infestation: yes (Hoogstraal et al. 1972)

Remarks: references to H. semermis published prior to Hoogstraal et al. (1965) have not been included in our analyses because of diagnostic uncertainties. Camicas et al. (1998) state that only ungulates are hosts of adults of this tick. We, however, believe that Hoogstraal et al. (1965) provide a reliable source of data indicating that hosts other than ungulates are also important for the natural maintenance of adults of H. semermis. We have been unable to find a description of the larva of H. semermis; nevertheless, we provisionally accept records of this stage in Hoogstraal et al. (1972). Lim (1972) recorded adults of H. semermis on Muridae, but Kolonin (2009) seems to have considered this record invalid and did not include Rodentia as hosts for adults. We regard Lim’s record as tentatively valid.

References

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. 1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.

Hoogstraal, H., Trapido, H. & Kohls, G.M. 1965. Southeast Asian Haemaphysalis ticks (Ixodoidea, Ixodidae). H. (Kaiseriana) papuana nadchatrami ssp. n. and redescription of H. (K.) semermis Neumann. J. Parasitol., 51: 433–451.

Hoogstraal, H., Lim, B.L., Nadchatram, M. & Anastos, G. 1972. The Gunong Benom Expedition 1967. 8. Ticks (Ixodidae) of Gunong Benom and their altitudinal distribution, hosts and medical relationships. Bull. Br. Mus. (Nat. Hist.) Zool., 23: 167–186.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Lim, B.L. 1972. Host-relationship and seasonal abundance of immature ticks (Haemaphysalis spp. and Dermacentor spp.) in primary and mixed-secondary rainforests in west Malaysia. Southeast Asia J. Trop. Med. Publ. Health, 3: 605–612.

138 – H. shimoga Trapido & Hoogstraal, 1964 (J. Parasitol., 50: 303–310)

Based on a comparison of descriptive illustrations, Kolonin (2009, op. cit. under H. anomaloceraea) treats H. shimoga as a synonym of H. taiwana, but we believe that illustrations provide insufficient grounds for such a decision. Camicas et al. (1998, op. cit. under H. anomaloceraea) regard H. anomaloceraea as a synonym of H. shimoga, without justifying this decision apart from a vague statement about “Kolonin, 1997 (in litt.),” a reference that we have been unable to find. See H. anomaloceraea.

Type depositories: USNTC (holotype, paratypes), BMNH, VRC, HH (paratypes) (Keirans and Hillyard 2001, op. cit. under H. aciculifer), as H. cornigera shimoga

Known stages: male, female, nymph, larva

Zoogeographic Region: Oriental

Ecoregions: tropical and subtropical dry broadleaf forests

Hosts: Rodentia: Muridae (A, N and/or L)

Artiodactyla: Bovidae, Cervidae, Suidae; Rodentia: Sciuridae (A)

Human infestation: yes (Tanskul and Inlao 1989)

Remarks: the larva and nymph of H. shimoga were described from laboratory-reared specimens (Trapido and Hoogstraal 1964). Tanskul and Inlao (1989) use the term “immatures” without specifying which of the immature stages naturally infested Muridae. These records are treated as provisionally valid here. See also H. cornigera.

References

Phan Trong, C. 1977. Ve bet va con trung ky sinh o Viet Nam. Tap 1. Ve (Ixodoidea), mo ta va phan loai. Ha Noi: Khoa hoc va ky thuat, 489 pp. In Vietnamese.

Rajagopalan, P.K., Patil, A.P. & Boshell, J. 1968. Ixodid ticks on their mammalian hosts in the Kyasanur Forest disease area of Mysore State, India, 1961–1964. Ind. J. Med. Res., 56: 510–526.

Tanskul, P. & Inlao, I. 1989. Keys to the adult ticks of Haemaphysalis Koch, 1844, in Thailand with notes on changes in taxonomy (Acari: Ixodoidea: Ixodidae). J. Med. Entomol., 26: 573–601.

Trapido, H. & Hoogstraal, H. 1964. Haemaphysalis cornigera shimoga subsp. n. from southern India (Ixodoidea, Ixodidae). J. Parasitol., 50: 303–310.

139 – H. silacea Robinson, 1912 (Parasitology, 4: 474–484)

Type depository: BMNH (lectotype, paralectotypes) (Keirans and Hillyard 2001, op. cit. under H. aciculifer)

Known stages: male, female, nymph, larva

Zoogeographic Region: Afrotropical

Ecoregions: tropical and subtropical grasslands, savannas and shrublands

Hosts: usual hosts for larvae, nymphs and adults are Artiodactyla: Bovidae.

Artiodactyla: Bovidae; Carnivora: Canidae; Lagomorpha: Leporidae (ANL)

Carnivora: Felidae (AN)

Carnivora: Herpestidae (A, N and/or L)

Macroscelidea: Macroscelididae; Galliformes: Numididae; Ciconiiformes: Threskiornithidae (NL)

Passeriformes: Alaudidae, Corvidae, Pycnonotidae (N)

Carnivora: Hyaenidae; Perissodactyla: Rhinocerotidae; Rodentia: Muridae (L)

Human infestation: yes (Horak et al. 2002)

Remarks: Theiler (1962) uses the term “immatures” without specifying whether they are larvae, nymphs or both. She also states that adults infest Aves but provides no data to support this observation.

References

Cumming, G.S. 1998. Host preference in African ticks (Acari: Ixodida): a quantitative data set. Bull. Entomol. Res., 88: 379–406.

Hoogstraal, H. 1963. Notes on African Haemaphysalis ticks. V. Redescription and relationships of H. silacea Robinson, 1912, from South Africa (Ixodoidea, Ixodidae). J. Parasitol., 49: 830–837.

Horak, I.G. 2001. Conservation of rare parasites. Proc. Int. Joint. Conf. Vet. Conserv. Biol. Wild Manag. Taronga Zoo, Sydney, Australia, Jul. 1–6, 2001, A. Martin & L. Vogelnest (editors), pp. 45–49.

Horak, I.G. & Knight, M.M. 1986. A comparison of the tick burdens of wild animals in a nature reserve and on an adjacent farm where tick control is practised. J. S. Afr. Vet. Ass., 57: 199–203.

Horak, I.G., Jacot Guillarmod, A., Moolman, L.C. & De Vos, V. 1987. Parasites of domestic and wild animals in South Africa. XXII. Ixodid ticks on domestic dogs and on wild carnivores. Onderstepoort J. Vet. Res., 54: 573–580.

Horak, I.G., Fourie, L.J., Heyne, H., Walker, J.B. & Needham, G.R. 2002. Ixodid ticks feeding on humans in South Africa: with notes on preferred hosts, geographic distribution, seasonal occurrence and transmission of pathogens. Exp. Appl. Acarol., 27: 113–136.

Horak, I.G., Welman, S., Hallam, S.L., Lutermann, H. & Mzilikazi, N. 2011. Ticks of four-toed elephant shrews and Southern African hedgehogs. Onderstepoort J. Vet. Res., 78, doi:10.4102/ojvr.v78i1.243

Knapp, S.E., Krecek, R.C., Horak, I.G. & Penzhorn, B.L. 1997. Helminths and arthropods of black and white rhinoceroses in southern Africa. J. Wildl. Dis., 33: 492–502.

Norval, R.A.I. 1975. Studies on the ecology of Haemaphysalis silacea Robinson 1912 (Acarina: Ixodidae). J. Parasitol., 61: 730–736.

Theiler, G. 1962. The Ixodoidea parasites of vertebrates in Africa south of the Sahara (Ethiopian Region). Report to the Director of Veterinary Services, Onderstepoort, South Africa, Project S.9958, 260 pp.

140 – H. silvafelis Hoogstraal & Trapido, 1963 (J. Parasitol., 49: 346–349)

Type depositories: USNTC (holotype, paratypes), ZSI, HH (paratypes) (Hoogstraal, H. & Trapido H. 1963. Haemaphysalis silvafelis sp. n., a parasite of the jungle cat in southern India (Ixodoidea: Ixodidae). J. Parasitol., 49: 346–349)

Known stages: male, female

Zoogeographic Region: Oriental

Ecoregion: eastern highlands moist deciduous forests

Hosts: usual hosts for adult ticks are Carnivora: Felidae.

Carnivora: Felidae; Lagomorpha: Leporidae (A)

Rodentia: Sciuridae; Cuculiformes: Cuculidae (N)

Human infestation: no

Remarks: Kolonin (2009) does not include Aves as hosts of H. silvafelis or recognize hosts for nymphs of this tick, probably because this stage remains undescribed. However, we regard the records of nymphs of H. silvafelis on Cuculidae and Sciuridae in Geevarghese and Dhanda (1995) as provisionally valid.

References

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. 1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.

Geevarghese, G. & Dhanda, V. 1995. Ixodid ticks of Maharashtra State, India. Acarologia, 36: 309–313.

Hoogstraal, H. & Trapido H. 1963. Haemaphysalis silvafelis sp. n., a parasite of the jungle cat in southern India (Ixodoidea: Ixodidae). J. Parasitol., 49: 346–349.

Kaul, H.N., Dhanda, V. & Mishra, A.C. 1979. A survey of ixodid ticks in Orissa State, India. Ind. J. Anim. Sci., 49: 707–712.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

141 – H. simplex Neumann, 1897 (Mém. Soc. Zool. Fr., 10: 324–420)

Type depository: BMNH (syntypes) (Keirans and Hillyard 2001, op. cit. under H. aciculifer)

Known stages: male, female, nymph, larva

Zoogeographic Region: Afrotropical

Ecoregion: Madagascar dry deciduous forests

Hosts: usual hosts for larvae, nymphs and adults are Afrosoricida: Tenrecidae. Aves and Primates are considered exceptional hosts for this tick.

Afrosoricida: Tenrecidae (ANL)

Primates: Indriidae (AN)

Rodentia: Muridae; Ciconiiformes: Threskiornithidae (A)

Primates: Cheirogaleidae (N)

Human infestation: no

Remarks: Camicas et al. (1998) list the larva of H. simplex as undescribed, but it had previously been described in Uilenberg et al. (1979). Primates are not included as hosts of H. simplex in Kolonin (2009), probably because Uilenberg et al. (1979) considered this type of host “accidental.” We see no reason to exclude Primates as hosts of this tick, although such parasitism may be unusual. The records of nymphs of H. simplex on Cheirogaleidae are considered provisionally valid by Rodríguez et al. (2012).

References

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. 1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Rodríguez, I.A., Rasoazanabary, E. & Godfrey, L.R. 2012. Multiple ectoparasites infest Microcebus griseorufus at Beza Mahafaly Special Reserve, Madagascar. Madagascar Cons. Develop., 7: 45–48.

Uilenberg, G., Hoogstraal, H. & Klein, J.-M. 1979. Les tiques (Ixodoidea) de Madagascar et leur rôle vecteur. Arch. Inst. Pasteur Madagascar Num. Spéc., 153 pp.

142 – H. simplicima Hoogstraal & Wassef, 1979 (In Uilenberg, G., Hoogstraal, H. & Klein, J.-M. 1979. Arch. Inst. Pasteur Madagascar Spec. Numb., 153 pp.)

Type depositories: USNTC (holotype, paratypes), BMNH, IPT (paratypes) (Keirans and Hillyard 2001, op. cit. under H. aciculifer)

Known stages: male, female, nymph

Zoogeographic Region: Afrotropical

Ecoregion: Madagascar spiny thickets

Hosts: Afrosoricida: Tenrecidae (AN)

Human infestation: no

Remarks: Durden and Keirans (1996) regard H. simplicima as an endangered species.

References

Durden, L.A. & Keirans, J.E. 1996. Host-parasite coextinction and the plight of tick conservation. Am. Entomol., 42: 87–91.

Uilenberg, G., Hoogstraal, H. & Klein, J.-M. 1979. Les tiques (Ixodoidea) de Madagascar et leur rôle vecteur. Arch. Inst. Pasteur Madagascar Num. Spéc., 153 pp.

143 – H. sinensis Zhang, 1981 (Acta Vet. Zoot. Sin., 12: 169–173. In Chinese)

This tick is ignored in Camicas et al. (1998, op. cit. under H. anomaloceraea), but it has been included in all recent lists of the ticks of the world. Consequently, we see no reason to question its validity.

Type depository: IAHVS (syntypes inferred by the English summary of the Chinese article) (Zhang, S.-X. 1981. A new species of Haemaphysalis Koch, 1844 – Haemaphysalis sinensis sp. nov. Acta Vet. Zoot. Sin., 12: 169–173. In Chinese)

Known stages: male, female

Zoogeographic Region: Palearctic

Ecoregions: montane grasslands and shrublands

Hosts: Artiodactyla: Bovidae (A)

Human infestation: no

References

Chen, Z., Yang, X., Bu, F., Yang, X., Yang, X. & Liu, J. 2010. Ticks (Acari: Ixodoidea: Argasidae, Ixodidae) of China. Exp. Appl. Acarol., 51: 393–404.

Keirans, J.E. & Robbins, R.G. 1999. A world checklist of genera, subgenera, and species of ticks (Acari: Ixodida) published from 1973 – 1997. J. Vector Ecol., 24: 115–129.

Zhang, S.-X. 1981. A new species of Haemaphysalis Koch, 1844 – Haemaphysalis sinensis sp. nov. Acta Vet. Zoot. Sin., 12: 169–173. In Chinese.

144 – H. spinigera Neumann, 1897 (Mém. Soc. Zool. Fr., 10: 324–420)

Type depository: ENV (holotype) (Trapido 1965, op. cit. under H. lagrangei)

Known stages: male, female, nymph, larva

Zoogeographic Region: Oriental

Ecoregions: tropical and subtropical dry broadleaf forests

Hosts: Mammals (several orders) (ANL)

Galliformes: Phasianidae (A?, NL)

Aves (several orders) (NL)

Human infestation: yes (Mitchell et al. 1966)

Remarks: Rajagopalan (1972) found adults of H. spinigera on Phasianidae, but Kolonin (2009) did not include Aves as hosts for adults of this tick, probably because Rajagopalan (1972) stated that adults found on Galliformes were teneral and apparently not feeding on these hosts.

References

Bhat, H.R. & Sreenivasan, M.A. 1981. Further records of the ticks of some reptilian and mammalian hosts in the Kyasanur Forest disease area, Karnataka, India. Ind. J. Parasitol., 5: 207–210.

Chen, Z., Yang, X., Bu, F., Yang, X., Yang, X. & Liu, J. 2010. Ticks (Acari: Ixodoidea: Argasidae, Ixodidae) of China. Exp. Appl. Acarol., 51: 393–404.

Kolonin, G.V. 1995. Review of the ixodid tick fauna (Acari: Ixodidae) of Vietnam. J. Med. Entomol., 32: 276–282.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Mitchell, R.M. & Dick, J.A. 1978. Ectoparasites from Nepal birds. J. Bombay Nat. Hist. Soc., 74: 264–274.

Mitchell, C.J., Hoogstraal, H., Schaller, G.B. & Spillett, J. 1966. Ectoparasites from mammals in Kanha National Park, Madhya Pradesh, India, and their potential disease relationships. J. Med. Entomol., 3: 113–124.

Rajagopalan, P.K. 1972. Ixodid ticks (Acarina: Ixodidae) parasitizing wild birds in the Kyasanur Forest disease area of Shimoga District, Mysore State, India. J. Bombay Nat. Hist. Soc., 69: 55–78.

Rajagopalan, P.K., Patil, A.P. & Boshell, J. 1968. Ixodid ticks on their mammalian hosts in the Kyasanur Forest disease area of Mysore State, India, 1961–1964. Ind. J. Med. Res., 56: 510–526.

Seneviratna, P. 1965. The Ixodoidea (ticks) of Ceylon. Parts II and III. Ceylon Vet. J., 13: 28–54.

Trapido, H., Goverdhan, M.K., Rajagopalan, P.K. & Rebello, M.J. 1964. Ticks ectoparasitic on monkeys in the Kyasanur Forest disease area of Shimoga District, Mysore State, India. Am. J. Trop. Med. Hyg., 13: 763–772.

145 – H. spinulosa Neumann, 1906 (Arch. Parasitol., 10: 195–219)

Type depository: BMNH (syntypes) (Neumann, L.G. 1906. Notes sur les Ixodidés. IV. Arch. Parasitol., 10: 195–219), but Keirans and Hillyard (2001, op. cit. under H. aciculifer) state that there are no types of H. spinulosa in BMNH.

Known stages: male, female, nymph, larva

Zoogeographic Region: Afrotropical

Ecoregions: tropical and subtropical grasslands, savannas and shrublands; few ticks in tropical and subtropical moist broadleaf forests

Hosts: usual hosts for adult ticks are Carnivora (several families).

Mammalia (several orders) (A)

Human infestation: no

Remarks: Horak et al. (1987) and Fourie et al. (1992) note the difficulties involved in distinguishing larvae and nymphs of H. spinulosa from those of H. elliptica (at the time identified as H. leachi). Consequently, references to these stages of H. spinulosa have been omitted, including those of Hussein and Mustafa (1985), who allegedly worked with larvae and nymphs of this species. Cumming (1998) includes Aves as hosts for H. spinulosa, but sound records for this tick on birds have not been found. Matthysse and Colbo (1987) state that the type specimen of H. spinulosa was collected from Rodentia: Thryonomyidae, but we have been unable to confirm this in Neumann (1906) or any subsequent publication. See also H. zumpti.

References

Cumming, G.S. 1998. Host preference in African ticks (Acari: Ixodida): a quantitative data set. Bull. Entomol. Res., 88: 379–406.

Fourie, L.J., Horak, I.G. & Van den Heever, J.J. 1992. The relative host status of rock elephant shrews Elephantulus myurus and Namaqua rock mice Aethomys namaquensis for economically important ticks. S. Afr. J. Zool., 27: 108–114.

Horak, I.G., Jacot Guillarmod, A., Moolman, L.C. & De Vos, V. 1987. Parasites of domestic and wild animals in South Africa. XXII. Ixodid ticks on domestic dogs and wild carnivores. Onderstepoort J. Vet. Res., 54: 573–580.

Hussein, H.S. & Mustafa, B.E. 1985. Haemaphysalis spinulosa and Rhipicephalus simus (Acari: Ixodidae): seasonal abundance of immature stages and host range in the Shambat area, Sudan. J. Med. Entomol., 22: 72–77.

Iori, A., Lanfranchi, P. & Manilla, G. 1996. Contribution to the knowledge of Ixodidae ticks of wild mammals of Somalia. Parassitologia, 38: 571–573.

Keirans, J.E. 1985. George Henry Falkiner Nuttall and the Nuttall tick catalogue. U. S. Dept. Agric., Agric. Res. Ser. Misc. Pub. (1438), 1785 pp.

Matthysse, J.G. & Colbo, M.H. 1987. The ixodid ticks of Uganda. Entomological Society of America, College Park, Maryland, 426 pp.

Morel, P.-C. & Rodhain, F. 1972. Contribution à la connaissance des tiques du sud de l’Éthiopie. Bull. Soc. Pathol. Exot., 65: 725–732.

Neumann, L.G. 1906. Notes sur les Ixodidés. IV. Arch. Parasitol., 10: 195–219.

Norval, R.A.I. 1984. The ticks of Zimbabwe. IX. Haemaphysalis leachi and Haemaphysalis spinulosa. Zimbabwe Vet. J., 15: 9–17.

146 – H. subelongata Hoogstraal, 1953 (Bull. Mus. Comp. Zool., 111: 37–113)

Type depositories: USNTC (holotype, paratypes), BMNH, FMNH, HH, MCZ, OVI, ISM (paratypes) (Keirans and Hillyard 2001, op. cit. under H. aciculifer)

Known stages: male, female, nymph, larva

Zoogeographic Region: Afrotropical

Ecoregion: Madagascar lowlands forests

Hosts: usual hosts for larvae, nymphs and adults are Afrosoricida-Tenricidae.

Afrosoricida: Tenricidae (ANL)

Rodentia: Muridae (N)

Rodentia: Nesomyidae (L)

Human infestation: no

Remarks: Durden and Keirans (1996) regard H. subelongata as an endangered species.

References

Durden, L.A. & Keirans, J.E. 1996. Host-parasite coextinction and the plight of tick conservation. Am. Entomol., 42: 87–91.

Uilenberg, G., Hoogstraal, H. & Klein, J.-M. 1979. Les tiques (Ixodoidea) de Madagascar et leur rôle vecteur. Arch. Inst. Pasteur Madagascar Num. Spéc., 153 pp.

147 – H. subterra Hoogstraal, El Kammah & Camicas, 1992 (Int. J. Acarol., 18: 213–220)

Type depositories: USNTC (holotype, paratypes), HH, ORSTOM (paratypes) (Hoogstraal, H., El Kammah, K.M. & Camicas, J.-L. 1992. Notes on African Haemaphysalis ticks: XVI. H. (Rhipistoma) subterra sp. n., a new member of the leachi group (Ixodoidea: Ixodidae). Int. J. Acarol., 18: 213–220)

Known stages: male, female, nymph, larva

Zoogeographic Region: Afrotropical

Ecoregions: tropical and subtropical grasslands, savannas and shrublands

Hosts: usual hosts for adult ticks are Carnivora: Herpestidae.

Rodentia: Spalacidae (ANL)

Carnivora: Herpestidae (AN)

Artiodactyla: Viverridae; Rodentia: Muridae (A)

Human infestation: no

Remarks: Hoogstraal et al. (1992) do not explicitly identify some hosts of the larvae of H. subterra. In the abstract of their paper, they state that “immatures” are found mainly on Rodentia (Muridae and Spalacidae), but in “material examined” there is no clear reference to larvae or nymphs of H. subterra feeding on Muridae. We thus provisionally exclude Muridae as hosts for the sub-adult stages of this tick.

Reference

Hoogstraal, H., El Kammah, K.M. & Camicas, J.-L. 1992. Notes on African Haemaphysalis ticks: XVI. H. (Rhipistoma) subterra sp. n., a new member of the leachi group (Ixodoidea: Ixodidae). Int. J. Acarol., 18: 213–220.

148 – H. sulcata Canestrini & Fanzago, 1878 (Atti R. Ist. Veneto Sci. Lett. Arti (1877–1878), Ser. 5, 4: 69–208)

See the initial section of this chapter for the synonymy of H. cholodkovskyi and H. cretica with H. sulcata.

Type depositories: undetermined

Known stages: male, female, nymph, larva

Zoogeographic Regions: Afrotropical, Oriental, Palearctic

Ecoregions: several ecoregions in the Afrotropical, Oriental and Palearctic Zoogeographic Regions

Hosts: usual hosts for adult ticks are Artiodactyla: Bovidae, while Testudines are considered exceptional hosts for adults and Chiroptera for larvae of this tick.

Testudines: Testudinidae (ANL)

Artiodactyla: Bovidae; Carnivora: Canidae (AN)

Several orders (Mammalia) (A)

Aves (several orders); Squamata (several families); Carnivora: Mustelidae; Lagomorpha: Ochotonidae; Rodentia: Cricetidae, Muridae (NL)

Chiroptera: Vespertilionidae; Rodentia: Calomyscidae, Dipodidae, Sciuridae (L)

Human infestation: yes (Bursali et al. 2012)

Remarks: Camicas et al. (1998) state that H. sulcata is found in the Oriental and Palearctic Regions, but Al-Khalifa et al. (2006) present sound records for the Afrotropical Region. Yeruham et al. (1996) refer to this tick as H. cretica.

References

Al-Khalifa, M.S., Diab, F.M., Al-Asgah, N.A., Hussein, H.S. & Khalil, G.M. 2006. Ticks (Acari: Argasidae, Ixodidae) recorded on wild animals in Saudi Arabia. Fauna Arabia, 22: 225–231.

Bursali, A., Keskin, A. & Tekin, S. 2012. A review of the ticks (Acari: Ixodida) of Turkey: species diversity, hosts and geographical distribution. Exp. Appl. Acarol., 57: 91–104.

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. 1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.

Chen, Z., Yang, X., Bu, F., Yang, X., Yang, X. & Liu, J. 2010. Ticks (Acari: Ixodoidea: Argasidae, Ixodidae) of China. Exp. Appl. Acarol., 51: 393–404.

Filippova, N.A. 1997. Ixodid ticks of subfamily Amblyomminae. Fauna of Russia and neighbouring countries, 4 (5), Nauka, St. Petersburg, 436 pp. In Russian.

Filippova, N.A., Neronov, V.M. & Farhang-Azad, A. 1976. Materials on the fauna of ixodids (Acarina, Ixodidae) of small mammals in Iran. Entomol. Obozr., 55: 467–478. In Russian.

Hoogstraal, H. & Kim, K.C. 1985. Tick and mammal coevolution, with emphasis on Haemaphysalis. In K.C. Kim (editor), Coevolution of parasitic arthropods and mammals. John Wiley & Sons, New York, pp. 505–568.

Keskin, A., Bursali, A., Kumlutas, Y., Ilgaz, C. & Tekin, S. 2013. The parasitism of immature stages of the Haemaphysalis sulcata (Acari: Ixodidae) on some reptiles in Turkey. J. Parasitol., in press.

Krčmar, S. 2012. Hard ticks (Acari, Ixodidae) of Croatia. ZooKeys, 234: 19–57.

Mihalca, A.D., Dumitrache, M.O., Magdaş, C., Gherman, C.M., Domşa, C., Mircean, V., Ghira, I.V., Pocora, V., Ionescu, D.T., Sikó Barabási, S., Cozma, V. & Sandor, A.D. 2012. Synopsis of the hard ticks (Acari: Ixodidae) of Romania with update on host associations and geographical distribution. Exp. Appl. Acarol., 58: 183–206.

Morel, P.-C. 2003. Les tiques d’Afrique et du Bassin méditerranéen (1965–1995). CIRAD- EMVT, 1342 pp.

Osipova, N.Z., Karas, F.R., Vargina, S.G. & Grebenyuk, Yu. I. 1975. Ectoparasites of wild animals in Crimean hemorrhagic fever natural focus of southern Kirgizia. In A.I. Protsenko (editor), Entomological investigations in Kirgizia. Akad. Nauk Kirgiz. SSR, Kirgiz. Otd. Vses. Entomol. Obshch. Izd., Frunze, pp. 124–125. In Russian, NAMRU-3 translation 1164.

Široký, P., Petrželková, K.J., Kamler, D., Mihalca, A.D. & Modrý, D. 2006. Hyalomma aegyptium as dominant tick in tortoises of the genus Testudo in Balkan countries, with notes on its host preferences. Exp. Appl. Acarol., 40: 279–290.

Yeruham, I., Hadani, A., Galker, F. & Rosen, S. 1996. The seasonal occurrence of ticks (Acari: Ixodidae) on sheep and in the field in the Judean area of Israel. Exp. Appl. Acarol., 20: 47–56.

149 – H. sumatraensis Hoogstraal, El Kammah, Kadarsan & Anastos, 1971 (J. Parasitol., 57: 1104–1109)

Type depositories: MZB (holotype, paratypes), USNTC, BMNH, HH (paratypes) (Hoogstraal, H., El Kammah, K.M., Kadarsan, S. & Anastos, G. 1971. Haemaphysalis (H.) sumatraensis sp. n. (Ixodoidea: Ixodidae), a tick parasitizing the tiger, boar and sambar deer in Indonesia. J. Parasitol., 57: 1104–1109), but Keirans and Hillyard (2001, op. cit. under H. aciculifer) state that there are no types ofH. sumatraensis in BMNH.

Known stages: male, female, larva

Zoogeographic Region: Oriental

Ecoregions: Sumatra and western Java rain forests

Hosts: Artiodactyla: Cervidae, Suidae; Carnivora: Canidae, Felidae (A)

Carnivora: Viverridae (N and/or L)

Human infestation: no

Remarks: Camicas et al. (1998) list the larva of H. sumatraensis as undescribed, but it was described by Kadarsan (1971). Hoogstraal and Kim (1985) use the term “immatures” without specifying whether they mean larvae, nymphs or both stages. The nymph of H. sumatraensis has not been described, but we consider the records of undetermined sub-adult stages in Hoogstraal and Kim (1985) provisionally valid.

References

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. 1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.

Hoogstraal, H. & Kim, K.C. 1985. Tick and mammal coevolution, with emphasis on Haemaphysalis. In K.C. Kim (editor), Coevolution of parasitic arthropods and mammals. John Wiley & Sons, New York, pp. 505–568.

Hoogstraal, H., El Kammah, K.M., Kadarsan, S. & Anastos, G. 1971. Haemaphysalis (H.) sumatraensis sp. n. (Ixodoidea: Ixodidae), a tick parasitizing the tiger, boar and sambar deer in Indonesia. J. Parasitol., 57: 1104–1109.

Kadarsan, S. 1971. Larval ixodid ticks of Indonesia (Acarina: Ixodidae). Ph.D. Dissertation, University of Maryland, 182 pp.

150 – H. sundrai Sharif, 1928 (Rec. Ind. Mus., 30: 217–344)

Ghosh, S., Bansal, G.C., Gupta, S.C., Ray, D., Khan, M.Q., Irshad, H., Shahiduzzaman, Md., Seitzer, U. & Ahmed, J.S. (2007. Status of tick distribution in Bangladesh, India and Pakistan. Parasitol. Res., 101 (Suppl. 2): 207–216) and Barker and Murrell (2008, op. cit. under H. colasbelcouri) treat Haemaphysalis himalaya Hoogstraal, 1966 and H. sundrai as valid species, but, as discussed in Guglielmone et al. (2009, op. cit. under H. colasbelcouri), the former is a synonym of the latter.

Type depository: USNTC (lectotype) (Keirans and Clifford 1984, op. cit. under H. bartelsi)

Known stages: male, female, nymph, larva

Zoogeographic Region: Oriental

Ecoregions: tropical and subtropical moist broadleaf forests

Hosts: Artiodactyla: Bovidae, Cervidae (ANL)

Human infestation: no

Remarks: Hoogstraal and El Kammah (1970) refer to this species as H. himalaya.

Reference

Hoogstraal, H. & El Kammah, K.M. 1970. Haemaphysalis (Herpetobia) himalaya Hoogstraal (Ixodoidea: Ixodidae), description of immature stages, hosts, and distribution. J. Parasitol., 56: 1023–1027.

151 – H. suntzovi Kolonin, 1993 (J. Med. Entomol., 30: 996–968)

Type depositories: ZIAC (holotype, paratypes), BMNH, MCZ, USNTC (paratypes) (Keirans and Hillyard 2001, op. cit. under H. aciculifer)

Known stages: male, female

Zoogeographic Region: Oriental

Ecoregion: northern Indochina subtropical forests

Hosts: Artiodactyla: Suidae; Rodentia: Hystricidae (A)

Human infestation: no

Reference

Kolonin, G.V. 1995. Review of the ixodid tick fauna (Acari: Ixodidae) of Vietnam. J. Med. Entomol., 32: 276–282.

152 – H. susphilippensis Hoogstraal, Kohls & Parrish, 1968 (J. Parasitol., 54: 616–621)

Type depositories: USNTC (holotype, paratypes), BMNH, FMNH, HH, BM (paratypes). (Hoogstraal, H., Kohls, G.M. & Parrish, D.W. 1968. Studies on Southeast Asian Haemaphysalis ticks (Ixodoidea, Ixodidae). H. (Kaiseriana) susphilippensis sp. n., a parasite of Luzon and Mindanao boars. J. Parasitol., 54: 616–621). Keirans and Hillyard (2001, op. cit. under H. aciculifer) report that there are no types of this species in BMNH.

Known stages: male, female

Zoogeographic Region: Oriental

Ecoregions: tropical and subtropical moist broadleaf forests

Hosts: usual hosts for adult ticks are Artiodactyla: Suidae.

Artiodactyla: Cervidae, Suidae (A)

Human infestation: no

Reference

Hoogstraal, H., Kohls, G.M. & Parrish, D.W. 1968. Studies on Southeast Asian Haemaphysalis ticks (Ixodoidea, Ixodidae). H. (Kaiseriana) susphilippensis sp. n., a parasite of Luzon and Mindanao boars. J. Parasitol., 54: 616–621.

153 – H. taiwana Sugimoto, 1936 (J. Soc. Trop. Agric. Taiwan, 8: 336–346. In Japanese)

Considerable confusion exists concerning this name. Teng, K.-F. & Jiang, Z.-J. (1991. Economic insect fauna of China. Fasc. 39, Acari: Ixodidae. Science Press, Beijing, 355 pp. In Chinese) regard H. taiwana as a synonym of H. cornigera, but Chen, Z., Yang, X., Bu, F., Yang, X., Yang, X. & Liu, J. (2010. Ticks (Acari: Ixodoidea: Argasidae, Ixodidae) of China. Exp. Appl. Acarol., 51: 393–404) list H. taiwana as present in China. Kolonin (2009, op. cit. under H. anomaloceraea) reduces H. shimoga to a synonym of H. taiwana. We consider H. taiwana a valid name, as discussed in Guglielmone et al. (2010, op. cit. under H. anomaloceraea), but it is obvious that a comparative study of these taxa is needed in order to solve this problem. See also H. anomaloceraea and H. shimoga.

Type depository: undetermined, as Haemaphysalis cornigera taiwana

Known stages: male, female

Zoogeographic Region: Oriental

Ecoregions: tropical and subtropical dry broadleaf forests

Hosts: Artiodactyla: Bovidae (A)

Human infestation: no

Remarks: Camicas et al. (1998) list the larva and nymph of H. taiwana as described, but we have been unable to find descriptions of either stage. Kolonin (2009) extends the host range of H. taiwana to include Cervidae and “small mammals,” probably because he includes hosts of H. shimoga among the hosts of H. taiwana. There are records of H. cornigera taiwana in the Chinese tick literature (i.e., Guo et al. 2000) that were not included in our analysis. See above.

References

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. 1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.

Guo, Y., Wan, K., Xu, S. et al. [sic] 2000. The discovery and study on the natural focus of Lyme disease in eastern Guandong Province. Chin. J. Zoon., 16: 42–45. In Chinese.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Luh, P.-L. & Woo, W.-C. 1950. A list of Chinese ticks. Chin. J. Entomol., 1: 195–222. In Chinese, NAMRU-3 translation 19.

154 – H. tauffliebi Morel, 1965 (Acarologia, 7: 281–285)

Type depository: not stated (Morel, P.-C. 1965. Description de Haemaphysalis tauffliebi n. sp. d’Afrique Central (Acariens, Ixodoidea). Acarologia, 7: 281–285)

Known stages: male, female

Zoogeographic Region: Afrotropical

Ecoregions: tropical and subtropical grasslands, savannas and shrublands

Hosts: Rodentia: Hystricidae (ANL)

Passeriformes: Motacillidae (AN)

Rodentia: Muridae; Carnivora: Felidae (N)

Human infestation: no

Remarks: Hoogstraal and Wassef (1973) state that the larva and nymph of H. tauffliebi are known, but we have been unable to find any description of them. Consequently, we regard records of larvae and nymphs on the hosts above as only provisionally valid. Cumming (1998) recognizes only Rodentia as hosts for this tick, but there are records in Morel and Rodhain (1972) from other orders of mammals, and Hoogstraal and Wassef (1973) and Hoogstraal and Kim (1985) list Aves as hosts for H. tauffliebi. Kolonin (2009) does not recognize Felidae, as reported by Morel and Rodhain (1972), or Aves as hosts.

References

Cumming, G.S. 1998. Host preference in African ticks (Acari: Ixodida): a quantitative data set. Bull. Entomol. Res., 88: 379–406.

Hoogstraal, H. & Kim, K.C. 1985. Tick and mammal coevolution, with emphasis on Haemaphysalis. In K.C. Kim (editor), Coevolution of parasitic arthropods and mammals. John Wiley & Sons, New York, pp. 505–568.

Hoogstraal, H. & Wassef, H.Y. 1973. The Haemaphysalis ticks (Ixodoidea: Ixodidae) of birds. 3.H. (Ornithophysalis) subgen. n.: definition, species, hosts and distribution in the Oriental, Palearctic, Malagasy, and Ethiopian Faunal Regions. J. Parasitol., 59: 1099–1117.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Morel, P.-C. & Rodhain, F. 1972. Contribution à la connaissance des tiques du sud de l’Éthiopie. Bull. Soc. Pathol. Exot., 65: 725–732.

Walker, J.B. 1991. A review of the ixodid ticks (Acari, Ixodidae) occurring in southern Africa. Onderstepoort J. Vet. Res., 58: 81–105.

155 – H. theilerae Hoogstraal, 1953 (Bull. Mus. Comp. Zool., 111: 37–113)

Type depositories: USNTC (holotype, paratypes), BMNH, FMNH, MCZ, MNHN, OVI (paratypes) (Hoogstraal, H. 1953. Ticks (Ixodoidea) of the Malagasy Faunal Region (excepting the Seychelles). Their origins and host-relationships; with descriptions of five new Haemaphysalis species. Bull. Mus. Comp. Zool., 111: 37–113), but Keirans and Hillyard (2001, op. cit. under H. aciculifer) state that there are no type specimens of H. theilerae in BMNH.

Known stages: male, female, nymph, larva

Zoogeographic Region: Afrotropical

Ecoregion: Madagascar lowland forests

Hosts: Afrosoricida: Tenrecidae (ANL)

Human infestation: no

Remarks: Durden and Keirans (1996) regard H. theilerae as an endangered species.

References

Durden, L.A. & Keirans, J.E. 1996. Host-parasite coextinction and the plight of tick conservation. Am. Entomol., 42: 87–91.

Uilenberg, G., Hoogstraal, H. & Klein, J.-M. 1979. Les tiques (Ixodoidea) de Madagascar et leur rôle vecteur. Arch. Inst. Pasteur Madagascar Num. Spéc., 153 pp.

156 – H. tibetensis Hoogstraal, 1965 (J. Parasitol., 51: 452–459)

Type depositories: BMNH (holotype, paratypes), USNTC (paratypes) (Keirans and Hillyard 2001, op. cit. under H. aciculifer)

Known stages: male, female, nymph

Zoogeographic Region: Palearctic

Ecoregions: montane grasslands and shrublands

Hosts: Carnivora: Canidae (AN)

Artiodactyla: Bovidae (A)

Human infestation: no

Remarks: Kolonin (2009) states that the immature stages of H. tibetensis have not been described, but the nymph had earlier been described by Hoogstraal (1965). Camicas et al. (1998) list only ungulates as hosts for adults of this tick, but this in error because the original description of H. tibetensis includes several specimens collected from Canidae.

References

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. 1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.

Chen, Z., Yang, X., Bu, F., Yang, X., Yang, X. & Liu, J. 2010. Ticks (Acari: Ixodoidea: Argasidae, Ixodidae) of China. Exp. Appl. Acarol., 51: 393–404.

Hoogstraal, H. 1965. Haemaphysalis tibetensis sp. n., and its significance in elucidating phylogenetic patterns in the genus (Ixodoidea, Ixodidae). J. Parasitol., 51: 452–459.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Teng, K.-F. 1982. Acarina: Ixodidae. In Insects of Xizang (Tibet) Volume II, Institute of Zoology, Academia Sinica, pp. 449–461. In Chinese, translation by Robbins, F.-M.Y.

157 – H. tiptoni Hoogstraal, 1953 (Bull. Mus. Comp. Zool., 111: 37–113)

Type depositories: USNTC (holotype, paratypes), BMNH, FMNH, HH, MCZ, OVI, MNHN, ISM (paratypes) (Keirans and Hillyard 2001, op. cit. under H. aciculifer)

Known stages: male, female

Zoogeographic Region: Afrotropical

Ecoregion: Madagascar lowland forests

Hosts: Afrosoricida: Tenrecidae (A)

Human infestation: no

Remarks: Durden and Keirans (1996) regard H. tiptoni as an endangered species.

References

Durden, L.A. & Keirans, J.E. 1996. Host-parasite coextinction and the plight of tick conservation. Am. Entomol., 42: 87–91.

Uilenberg, G., Hoogstraal, H. & Klein, J.-M. 1979. Les tiques (Ixodoidea) de Madagascar et leur rôle vecteur. Arch. Inst. Pasteur Madagascar Num. Spéc., 153 pp.

158 – H. toxopei Warburton, 1927 (Parasitology, 19: 405–410)

Kolonin (2009, op. cit. under H. anomaloceraea) excludes H. toxopei from his list of ixodids of the world, giving no reason for doing so. Perhaps he followed Anastos (1950, op. cit. under H. hylobatis), who regards H. toxopei as a synonym of H. kinneari based on Warburton, C. (1927. On five new species of ticks (Arachnida Ixodoidea), Ornithodorus [sic] nattereri, Ixodes theodori, Haemaphysalis toxopei, Amblyomma robinsoni and A. dammermani, with a note on the ornate nymph of A. latum. Parasitology, 19: 405–410). However, Warburton’s statement is treated as a lapsus in Trapido, H., Hoogstraal, H. & Varma, M.G.R. (1964. Status and descriptions of Haemaphysalis p. papuana Thorell (n. comb.) and of H. papuana kinneari Warburton (n. comb.) (Ixodidae) of southern Asia and New Guinea. J. Parasitol., 50: 172–188). Haemaphysalis toxopei is a valid species.

Type depository: BMNH (lectotype, paralectotypes) (Keirans and Hillyard 2001, op. cit. under H. aciculifer)

Known stages: male, female

Zoogeographic Regions: Australasian, Oriental

Ecoregions: tropical and subtropical moist broadleaf forests

Hosts: Chiroptera: Pteropodidae (AN)

Carnivora: Canidae (A)

Human infestation: no

Remarks: although the nymphal stage of H. toxopei has not been described, we regard a single collection from Chiroptera in Durden et al. (2008) as provisionally valid. Hoogstraal (1964) refers to this tick as H. papuana toxopei.

References

Durden, L.A., Merker, S. & Beati, L. 2008. The tick fauna of Sulawesi, Indonesia (Acari: Ixodoidea: Argasidae and Ixodidae). Exp. App. Acarol., 45: 85–110.

Hoogstraal, H. 1964. Studies on Southeast Asian Haemaphysalis ticks (Ixodoidea, Ixodidae). Redescription, hosts, and distribution of H. traguli Oudemans. The larva and nymph of H. vidua W. and N. Identity of H. papuana toxopei Warburton (n. comb.). J. Parasitol., 50: 765–782.

159 – H. traguli Oudemans, 1928 (Entomol. Ber., 7 (164): 374–383)

Type depository: LMNH (syntypes) (Hoogstraal, H. 1964. Studies on Southeast Asian Haemaphysalis ticks (Ixodoidea, Ixodidae). Redescription, hosts, and distribution of H. traguli Oudemans. The larva and nymph of H. vidua W. and N. Identity of H. papuana toxopei Warburton. J. Parasitol., 50: 765–782)

Known stages: male, female, nymph, larva

Zoogeographic Region: Oriental

Ecoregions: tropical and subtropical moist broadleaf forests; few ticks in tropical and subtropical dry broadleaf forests

Hosts: usual hosts for larvae, nymphs and adults are Artiodactyla: Tragulidae.

Artiodactyla: Tragulidae (ANL)

Artiodactyla: Suidae; Rodentia: Muridae (A)

Human infestation: no

Remarks: Nadchatram et al. (1966) regard diagnoses of nymphs of H. traguli from Scandentia as tentative, and these hosts have therefore not been included in our list. These authors also erroneously refer to H. traguli as H. atheruri on page 133 of their paper. Kolonin (2009) excludes Suidae from his list of hosts for this tick, but we regard the record from this type of host in Keirans (1985) as valid.

References

Hoogstraal, H. 1964. Studies on Southeast Asian Haemaphysalis ticks (Ixodoidea, Ixodidae). Redescription, hosts, and distribution of H. traguli Oudemans. The larva and nymph of H. vidua W. and N. Identity of H. papuana toxopei Warburton. J. Parasitol., 50: 765–782.

Keirans, J.E. 1985. George Henry Falkiner Nuttall and the Nuttall tick catalogue. U. S. Dept. Agric., Agric. Res. Ser. Misc. Pub. (1438), 1785 pp.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Nadchatram, M., Domrow, R. & Ng, C.K. 1966. Parasitic Acarina of the mammals. Bull. Nat. Mus. State Singapore (34): 129–140.

160 – H. traubi Kohls, 1955 (J. Parasitol., 41: 312–315)

Type depositories: USNTC (holotype, paratype), BMNH (paratype) (Keirans & Hillyard, 2001, op. cit. under H. aciculifer).

Known stages: male, female

Zoogeographic Region: Oriental

Ecoregion: Borneo rain forests

Hosts: Artiodactyla: Cervidae, Suidae (A)

Human infestation: no

Reference

Hoogstraal, H. & El Kammah, K.M. 1971. Studies on Southeast Asian Haemaphysalis ticks (Ixodoidea: Ixodidae). H. (H.) traubi Kohls, redescription of male, description of female, and new artiodactyl host and Malayan distribution records. J. Parasitol., 57: 426–431.

161 – H. turturis Nuttall & Warburton, 1915 (Ticks. A monograph of the Ixodoidea. Part III. The genus Haemaphysalis. Cambridge University Press, London, pp. 349–550)

Type depository: ZMB (holotype) (Moritz and Fischer 1981, op. cit. under H. cinnabarina)

Known stages: male, female, nymph, larva

Zoogeographic Region: Oriental

Ecoregions: tropical and subtropical dry broadleaf forests

Hosts: Aves are considered exceptional hosts for adults of this tick.

Mammalia (several orders); Cuculiformes: Cuculidae; Galliformes: Phasianidae; Passeriformes: Turdidae (ANL)

Columbiformes: Columbidae (A)

Passeriformes (several families); Piciformes: Picidae (NL)

Psittaciformes: Psittacidae (N)

Human infestation: yes (Prakasan and Ramani 2003)

Remarks: Camicas et al. (1998) unaccountably do not specify any hosts for the larvae and nymphs of H. turturis. Kolonin (2009) ignores Aves as hosts for adult H. turturis, but there is evidence for this host-parasite relationship in Trapido et al. (1963), who redescribed the holotype collected from Columbidae, and in Rajagopalan (1972), who found small numbers of adult H. turturis on Aves.

References

Bhat, H.R. & Sreenivasan, M.A. 1981. Further records of the ticks of some reptilian and mammalian hosts in the Kyasanur Forest disease area, Karnataka, India. Ind. J. Parasitol., 5: 207–210.

Boshell, J. & Rajagopalan, P.K. 1968. Observations on the experimental exposure of monkeys, rodents and shrews to infestation of ticks in forest in Kyasanur Forest disease area. In. J. Med. Res., 56 (4) (Suppl.): 573–588.

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. 1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.

Hoogstraal, H. & Rack, G. 1967. Ticks (Ixodidae) collected by Deutsche Indien-Expedition, 1955–1958. J. Med. Entomol., 3: 284–288.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Prakasan, K. & Ramani, M. 2003. Human infesting ixodid ticks of Kerala. J. Parasit. Dis., 27: 108–112.

Rajagopalan, P.K. 1972. Ixodid ticks (Acarina: Ixodidae) parasitizing wild birds in the Kyasanur Forest disease area of Shimoga District, Mysore State, India. J. Bombay Nat. Hist. Soc., 69: 55–78.

Rajagopalan, P.K. & Sreenivasan, M.A. 1981. Ixodid ticks on cattle and buffaloes in the Kyasanur Forest disease area of Karnataka State. Ind. J. Med. Res., 73: 880–889.

Rajagopalan, P.K., Patil, A.P. & Boshell, J. 1968. Ixodid ticks on their mammalian hosts in the Kyasanur Forest disease area of Mysore State, India, 1961–1964. Ind. J. Med. Res., 56: 510–526.

Trapido, H., Hoogstraal, H. & Varma, M.G.R. 1963. Redescription of the holotype and description of all stages of Haemaphysalis turturis Nuttall and Warburton from Ceylon and India (Ixodoidea, Ixodidae). J. Parasitol., 49: 678–685.

Trapido, H., Goverdhan, M.K., Rajagopalan, P.K. & Rebello, M.J. 1964. Ticks ectoparasitic on monkeys in the Kyasanur Forest disease area of Shimoga District, Mysore State, India. Am. J. Trop. Med. Hyg., 13: 763–772.

162 – H. verticalis Itagaki, Noda & Yamaguchi, 1944 (In Ticks parasitic on domestic animals of Asia. Japanese Horse-Racing Association, Tokyo, 97 pp. In Japanese)

Type depository: NSM (syntypes, but apparently lost) (Emel’yanova, N.D. & Hoogstraal, H. 1973. Haemaphysalis verticalis Itagaki, Noda, and Yamaguchi: rediscovery in China, adult and immature identity, rodent hosts, distribution, and medical relationships (Ixodoidea: Ixodidae). J. Parasitol., 59: 724–733)

Known stages: male, female, nymph, larva

Zoogeographic Region: Palearctic

Ecoregions: temperate grasslands, savannas and shrublands

Hosts: usual hosts for larvae, nymphs and adults are Rodentia: Sciuridae. Aves are considered exceptional hosts for this tick.

Rodentia: Sciuridae, Muridae (ANL)

Mammalia (several orders); Passeriformes: Passeridae (A)

Human infestation: no

Remarks: the information obtained on host relationships in Teng and Jiang (1991) and Ma (1996) is inferred, and because of our inability to fully understand what stages were found on particular hosts, we have limited ourselves to adult ticks. Kolonin (2009) excludes Aves as hosts for H. verticalis, but we consider a record in Teng and Jiang (1991) from Passeriformes as provisionally valid.

References

Chen, Z., Yang, X., Bu, F., Yang, X., Yang, X. & Liu, J. 2010. Ticks (Acari: Ixodoidea: Argasidae, Ixodidae) of China. Exp. Appl. Acarol., 51: 393–404.

Emel’yanova, N.D. & Hoogstraal, H. 1973. Haemaphysalis verticalis Itagaki, Noda, and Yamaguchi: rediscovery in China, adult and immature identity, rodent hosts, distribution, and medical relationships (Ixodoidea: Ixodidae). J. Parasitol., 59: 724–733.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Ma, L. 1996. Observation on the behaviour of Haemaphysalis verticalis in the meadow of Jilin Province. Acta Entomol. Sin., 39: 219–222. In Chinese.

Teng, K.-F. & Jiang, Z.-J. 1991. Economic insect fauna of China. Fasc. 39, Acari: Ixodidae. Science Press, Beijing, 355 pp. In Chinese.

Zhuo, H.-M. & Su, J.-Y. 2003. Study of Lyme disease in Xilinguole District of Inner Mongolia. Chin. J. Vet. Med., 39: 6–7. In Chinese.

163 – H. vidua Warburton & Nuttall, 1909 (Parasitology, 2: 57–76)

Type depository: BMNH (holotype) (Keirans & Hillyard, 2001, op. cit. under H. aciculifer)

Known stages: male, female, nymph, larva

Zoogeographic Region: Oriental

Ecoregions: tropical and subtropical moist broadleaf forests

Hosts: Carnivora: Viverridae (ANL)

Human infestation: no

Reference

Hoogstraal, H. 1964. Studies on Southeast Asian Haemaphysalis ticks (Ixodoidea, Ixodidae). Redescription, hosts, and distribution of H. traguli Oudemans. The larva and nymph of H. vidua W. and N. Identity of H. papuana toxopei Warburton (n. comb.). J. Parasitol., 50: 765–782.

164 – H. warburtoni Nuttall, 1912 (Parasitology, 5: 50–60)

Type depository: BMNH (lectotype, paralectotypes) (Keirans and Hillyard 2001, op. cit. under H. aciculifer)

Known stages: male, female, nymph, larva

Zoogeographic Regions: Oriental, Palearctic

Ecoregion: lower Gangetic Plains moist deciduous forests

Hosts: Artiodactyla: Bovidae (ANL)

Rodentia: Muridae; Galliformes: Phasianidae (NL)

Human infestation: no

Remarks: papers on H. warburtoni adults published prior to Hoogstraal (1966) and those on larvae and nymphs published prior to Hoogstraal (1971) have not been considered in our analyses because of diagnostic uncertainties. Mitchell (1979) listed all the hosts of Nepalese ticks collected from 1966 to 1970, including those of H. warburtoni (no developmental stage provided), which was allegedly found on Cricetidae. This host is not included in Hoogstraal (1971) and it has been excluded from our list for H. warburtoni. There is also a record of H. warburtoni feeding on a human in Kirghizia (Sartbaev 1955), but according to the range statement in Hoogstraal (1966), the species was probably H. pospelovashtromae. Pending resolution of this matter, we have excluded humans from our list of hosts of H. warburtoni and H. pospelovashtromae.

References

Hoogstraal, H. 1966. Haemaphysalis (Allophysalis) pospelovashtromae sp. n. from USSR and redescription of the type material of H. (A.) warburtoni Nuttall from China (Ixodoidea, Ixodidae). J. Parasitol., 52: 787–800.

Hoogstraal, H. 1971. Haemaphysalis (Allophysalis) warburtoni Nuttall: description of immature stages, adult structural variation, and hosts and ecology in Nepal, with a redefinition of the subgenus Allophysalis Hoogstraal (Ixodoidea: Ixodidae). J. Parasitol., 57: 1083–1095.

Mitchell, R.M. 1979. A list of ectoparasites from Nepalese mammals, collected during the Nepal ectoparasite program. J. Med. Entomol., 16: 227–233.

Sartbaev, S.K. 1955. Materials on the biology of Haemaphysalis warburtoni Nutt., 1912, in conditions of Kirghizia (preliminary information). Trudy Inst. Zool. Parasitol. Akad. Nauk Kirgiz, SSR (4): 121–127. In Russian, NAMRU-3 translation 12.

165 – H. wellingtoni Nuttall & Warburton, 1908 (Proc. Cambr. Philos. Soc., 14: 392–416)

Type depository: BMNH (lectotype, paralectotypes) (Keirans & Hillyard, 2001, op. cit. under H. aciculifer)

Known stages: male, female, nymph, larva

Zoogeographic Regions: Australasian, Oriental

Ecoregions: tropical and subtropical moist broadleaf forests; few ticks in tropical and subtropical dry broadleaf forests

Hosts: usual hosts for larvae, nymphs and adults are Galliformes: Phasianidae.

Mammalia (several orders); Aves (several orders) (ANL)

Human infestation: yes (Durden et al. 2008)

Remarks: Yamaguti et al. (1971) and Miyamoto et al. (1993) provide bona fide records of H. wellingtoni for the Palearctic Region, but the ticks had been found on migratory birds and this species is probably not established there. Joyce (1965) reports its introduction into remote Pacific islands, where it has also failed to become established.

References

Bhat, H.R. & Sreenivasan, M.A. 1981. Further records of the ticks of some reptilian and mammalian hosts in the Kyasanur Forest disease area, Karnataka, India. Ind. J. Parasitol., 5: 207–210.

Boshell, J. & Rajagopalan, P.K. 1968. Observations on the experimental exposure of monkeys, rodents and shrews to infestation of ticks in forest in Kyasanur Forest disease area. Ind. J. Med. Res., 56 (4) (Suppl.): 573–588.

Durden, L.A., Merker, S. & Beati, L. 2008. The tick fauna of Sulawesi, Indonesia (Acari: Ixodoidea: Argasidae and Ixodidae). Exp. App. Acarol., 45: 85–110.

Emel’yanova, N.D. & Shikhorbeev, B.V. 1971. Haemaphysalis wellingtoni Nutt. and W. 1907 – a new ixodid tick in the USSR fauna. Dokl. Irkutsk. Protiv. Inst. (9): 249. In Russian, NAMRU-3 translation 574.

Hoogstraal, H., Lim, B.L., Nadchatram, M. & Anastos, G. 1972. The Gunong Benom Expedition 1967. 8. Ticks (Ixodidae) of Gunong Benom and their altitudinal distribution, hosts and medical relationships. Bull. Br. Mus. (Nat. Hist.) Zool., 23: 167–186.

Joyce, C.R. 1965. Notes and exhibitions. Proc. Hawaiian Entomol. Soc., 19: 25.

Miyamoto, K., Nakao, M, Fujita, H. & Sato, F. 1993. The ixodid ticks on migratory birds in Japan and the isolation of Lyme disease spirochetes from bird-feeding ticks. Jap. J. Sanit. Zool., 44: 315–326.

Parola, P., Cornet, J.-P., Sanogo, Y.O., Miller, R.S., Thien, H.V., González, J.-P., Raoult, D., Telford, S.R. & Wongsrichanalai, C. 2003. Detection of Ehrlichia spp., Anaplasma spp., Rickettsia spp., and other eubacteria in ticks from the Thai-Myanmar border and Vietnam. J. Clin. Microbiol., 41: 1600–1608.

Rajagopalan, P.K. 1972. Ixodid ticks (Acarina: Ixodidae) parasitizing wild birds in the Kyasanur Forest disease area of Shimoga District, Mysore State, India. J. Bombay Nat. Hist. Soc., 69: 55–78.

Tanskul, P., Stark, H.E. & Inlao, I. 1983. A checklist of ticks of Thailand (Acari: Metastigmata: Ixodoidea). J. Med. Entomol., 20: 330–341.

Trapido, H., Goverdhan, M.K., Rajagopalan, P.K. & Rebello, M.J. 1964. Ticks ectoparasitic on monkeys in the Kyasanur Forest disease area of Shimoga District, Mysore State, India. Am. J. Trop. Med. Hyg., 13: 763–772.

Xu, R. & Li, K. 1997. A collection of ticks from Guizhou, China. Syst. Appl. Acarol., 2: 245–246.

Yamaguti, N., Tipton, V.J., Keegan, H.L. & Toshioka, S. 1971. Ticks of Japan, Korea, and the Ryukyu Islands. Brigham Young Univ. Sci. Bull. Biol. Ser., 15 (1), 226 pp.

166 – H. yeni Toumanoff, 1944 (In Les tiques (Ixodoidea) de l’Indochine. Instituts Pasteur de l’Indochine, S.I.L.I., Saigon, 220 pp.)

Type depository: IPP (lectotype) (Hoogstraal, H. & Trapido, H. 1966. Redescription of the type materials of Haemaphysalis (Kaiseriana) bispinosa Neumann (India), H. (K.) neumanni Dönitz (Japan), H. (K.) lagrangei Larrousse (Vietnam), and H. (K.) yeni Toumanoff (Vietnam) (Ixodoidea: Ixodidae). J. Parasitol., 52: 1188–1198)

Known stages: male, female, nymph, larva

Zoogeographic Regions: Oriental, Palearctic

Ecoregions: tropical and subtropical dry broadleaf forests

Hosts: usual hosts for adult ticks are Artiodactyla: Cervidae.

Artiodactyla: Cervidae; Lagomorpha: Leporidae (ANL)

Carnivora: Canidae (AN)

Carnivora: Felidae, Viverridae (A, N and/or L)

Artiodactyla: Bovidae (A)

Human infestation: no

Remarks: Camicas et al. (1998) state that this tick is found exclusively in the Oriental Zoogeographic Region, but there are now also bona fide records of H. yeni for the Palearctic Region in Inokuma et al. (2002), among others. Kolonin (2009) mentions immature stages feeding on several hosts without specifying whether larvae, nymphs or both stages of H. yeni were found on Viverridae. He also ignores Leporidae as hosts for this tick, but we accept records from this type of host in Xu and Luo (1998).

References

Camicas, J.-L., Hervy, J.-P., Adam, F. & Morel, P.-C. 1998. Les tiques du monde (Acarida, Ixodida). Nomenclature, stades décrits, hôtes, répartition. ORSTOM, Paris, 233 pp.

Cao, W.-C., Gao, Y.-M., Zhang, P.-H., Zhang, X.-T., Dai, Q.-H., Dumler, J.S., Fang, L.-Q. & Yang, H. 2000. Identification of Ehrlichia chaffensis by nested PCR in ticks from southern China. J. Clin. Microbiol., 38: 2778–2880.

Chen, Z., Yang, X., Bu, F., Yang, X., Yang, X. & Liu, J. 2010. Ticks (Acari: Ixodoidea: Argasidae, Ixodidae) of China. Exp. Appl. Acarol., 51: 393–404.

Inokuma, H., Fujimoto, T., Hoso, E., Tanaka, S., Fujisaki, K., Okuda, M. & Onishi, T. 2002. Tick infestation of sika deer (Cervus nippon) in the western part of Yamaguchi Prefecture, Japan. J. Vet. Med. Sci., 64: 615–617.

Kolonin, G.V. 1995. Review of the ixodid tick fauna (Acari: Ixodidae) of Vietnam. J. Med. Entomol., 32: 276–282.

Kolonin, G.V. 2009. Fauna of ixodid ticks of the world. http://www.kolonin.org/

Phan Trong, C. 1977. Ve bet va con trung ky sinh o Viet Nam. Tap 1. Ve (Ixodoidea), mo ta va phan loai. Ha Noi: Khoa hoc va ky thuat, 489 pp. In Vietnamese.

Saito, Y. & Hoogstraal, H. 1972. Haemaphysalis (Kaiseriana) yeni Toumanoff (Ixodoidea: Ixodidae): discovery in Japan, description of female and immature stages, environment and life cycle. J. Parasitol., 58: 950–959.

Xu, R. & Luo, G. 1998. Ticks parasitic on Muntiacus reevesi and Lepus sinensis in Wuyi Mountains, Fujian, China. Syst. Appl. Acarol., 3: 197.

167 – H. zumpti Hoogstraal & El Kammah, 1974 (J. Parasitol., 60: 188–197)

Type depositories: USNTC (holotype, paratypes), BMNH, FMNH, HH, OVI, SAIMR, CM (paratypes) (Keirans and Hillyard 2001, op. cit. under H. aciculifer)

Known stages: male, female, nymph, larva

Zoogeographic Region: Afrotropical

Ecoregion: southern Africa bushveld

Hosts: Carnivora: Herpestidae, Hyaenidae; Rodentia: Sciuridae (ANL)

Carnivora: Viverridae (AN)

Carnivora (several orders) (A)

Human infestation: no

Remarks: Cumming (1998) states that Carnivora are the only hosts for H. zumpti, but this species has also been collected from Sciuridae (Hoogstraal and El Kammah 1974). Walker (1991) stresses the difficulties involved in separating larvae and nymphs of H. zumpti from those of H. leachi and H. spinulosa. However, we accept the records in Hoogstraal and El Kammah (1974) and Horak et al. (2000) of larval and nymphal H. zumpti on the hosts listed above.

References

Cumming, G.S. 1998. Host preference in African ticks (Acari: Ixodida): a quantitative data set. Bull. Entomol. Res., 88: 379–406.

Hoogstraal, H. & El Kammah, K.M. 1974. Notes on African Haemaphysalis ticks. XII. H. (Rhipistoma) zumpti sp. n., a parasite of small carnivores and squirrels in southern Africa (Ixodoidea: Ixodidae). J. Parasitol., 60: 188–197.

Horak, I.G., Braack, L.E.O., Fourie, L.J. & Walker, J.B. 2000. Parasites of domestic and wild animals in South Africa. XXXVIII. Ixodid ticks collected from 23 wild carnivore species. Onderstepoort J. Vet. Res., 67: 239–250.

Matthee, S., Lovely, C., Gaugler, A., Beeker, R., Venter, H.R. & Horak, I.G. 2010. Ixodid ticks on domestic dogs in the Northern Cape Province of South Africa and in Namibia. J. S. Afr. Vet. Assoc., 81: 126–128.

Walker, J.B. 1991. A review of the ixodid ticks (Acari, Ixodidae) occurring in southern Africa. Onderstepoort J. Vet. Res., 58: 81–105.

Synopsis of the Genus Haemaphysalis

A synopsis of the genus Haemaphysalis is presented in Tables 1 and 2. The Oriental Region contains the greatest number of species occurring exclusively in a single zoogeographic region (64, or 38 % of the total), while the Neotropical and Nearctic Regions contain the least number of species confined to a single region, namely H. cinnabarina (known from two specimens collected almost 170 years ago) and H. chordeilis, respectively. Six species are widely distributed. Four of these species – H. cornigera, H. doenitzi, H. hystricis and H. longicornis – occur in the Australasian, Oriental and Palearctic Zoogeographic Regions, while two species – H. indica and H. sulcata – occur in the Afrotropical, Oriental and Palearctic Regions. As a genus, haemaphysalids are confined to 12 combinations of zoogeographic regions.

Table 1 Numbers and percentages of all species of Haemaphysalis, by zoogeographic region(s), number known to feed on humans (% of the total number of species in a particular region) and number of species for which all stages (larva, nymph, male and/or female) are known (% of the total number of species in a particular region)

Table 2 Number and percentages of the 85 species of Haemaphysalis whose adult (female and/or male), larval and nymphal stages as well as natural hosts are known, including or excluding exceptional hosts

Our overall analyses indicate that 118 species (71 %) occur in regions that constituted Gondwanaland, 20 species (12 %) are found in lands that formed Laurasia, and the remaining 28 species (17 %) occur in areas that were part of both ancient continents.

A total of 56 species (33 %) have been recorded as feeding on humans.

The larva, nymph and at least one adult stage are known for just 98 (59 %) of the 167 species of Haemaphysalis that we recognize as valid. Clearly, efforts should be made to expand our knowledge of the taxonomy of the subadult stages of this genus in order to better understand the phylogenetic relationships of haemaphysalids as a whole. Nevertheless, most parasitic stages of species within the Palearctic and Oriental-Palearctic Region are known.

Host utilization by the 85 species of Haemaphysalis whose adult (female and/or male) and sub-adult stages are known, together with their natural hosts, is presented in Table 2, first including exceptional hosts, and then excluding these hosts. Although all parasitic stages of H. colasbelcouri, H. colesbergensis, H. fujisana, H. kitaokai, H. megaspinosa, H. norvali, and H. shimoga are known, one or more of these stages are known only from laboratory-reared specimens. On the other hand, the natural hosts of the larva or the nymph of H. indoflava, H. leachi, H. montgomeryi, H. obesa and H. spinulosa are not known with certainty, and while the male, female, nymph and larva of H. kumaonensis are known, their hosts are not. These 13 species have therefore been excluded from our analysis.

When exceptional hosts are included, Haemaphysalis ticks parasitize seven categories of hosts (Table 2). The only exclusive hosts are Mammalia (29 species or 34 % of the total), and Aves, which are exclusive hosts for the Oriental H. megalaimae. However, Mammalia are exclusive and non-exclusive hosts for 84 species (99 %), and Aves are exclusive or non-exclusive hosts for 55 species (65 %). The most common combination of hosts is Mammalia + Aves (45 species or 53 %). By comparison, the contribution of Squamata and Testudines is insignificant because these groups are hosts of just 8 and 6 species of Haemaphysalis, respectively, and even then only in combination with two to three other types of hosts for the corresponding species (Table 2). Anura and Crocodilia are not known to host Haemaphysalis ticks.

The pattern of host utilization undergoes important changes when exceptional hosts are excluded from the analysis. First, the categories of hosts diminish to six, because the combination Aves + Mammalia + Testudines drops out. Mammalia increases its share as exclusive hosts for almost half of the species (41, or 48 %). Four species – H. chordeilis, H. doenitzi, H. hoodi and H. minuta – join H. megalaimae as exclusive parasites of Aves (6 % of the total). The most common combination of hosts is still Mammalia + Aves (34 species, or 40 %), followed by the three combinations of hosts shown in Table 2. The representation of Squamata as hosts is now reduced to five species, the Palearctic H. caucasica, H. parva and H. punctata, the Oriental H. kashmirensis, and the Afrotropical, Oriental and Palearctic species H. sulcata. However, the role of Squamata is certainly relevant for the maintenance of larvae and nymphs of H. kashmirensis. Haemaphysalis sulcata is the only species with Testudinidae as non-exclusive hosts when exceptional hosts are excluded from the analysis; among other types of hosts, the larvae and nymphs of H. sulcata are also found on Testudinidae and on Squamata.