Experimental and Applied Acarology 28: 2754, 2002.

2003 Kluwer Academic Publishers. Printed in the Netherlands.

The Argasidae, Ixodidae and Nuttalliellidae (Acari:

Ixodida): a world list of valid tick names

IVAN G. HORAK1,, JEAN-LOUIS CAMICAS2

and JAMES E. KEIRANS3
1 Department of Veterinary Tropical Diseases, Faculty of Veterinary Science, University of
Pretoria, Onderstepoort, 0110 South Africa
2 Laboratoire dAcarologie mdicale, Centre IRD de Montpellier, BP 64501, 34394
Montpellier Cedex 5, France
3 Curator United States National Tick Collection, Georgia Southern University, Statesboro,
GA 30460-8056, USA

Abstract. The worlds argasid tick fauna comprises 183 species in four genera, namely Argas,
Carios, Ornithodoros and Otobius in the family Argasidae. The ixodid tick fauna consists of
241 species in the genus Ixodes and 442 species in the genera Amblyomma, Anomalohima-
laya, Bothriocroton, Cosmiomma, Dermacentor, Haemaphysalis, Hyalomma, Margaropus,
Nosomma, Rhipicentor and Rhipicephalus in the family Ixodidae, with the genus Boophilus
becoming a subgenus of the genus Rhipicephalus. The family Nuttalliellidae is represented by
the monospecific genus Nuttalliella. The species names of these ticks, based on seven previous
complete or partial listings, as well as those of recently described new species, are presented
in tabular format.

Key words: ticks, Argasidae, Ixodidae, Nuttalliellidae, world checklist, species names

Preamble

We believe that the animal known as the species exists in the world, and that
it can be given a name using binominal nomenclature. Only the species exists
in nature, and any organization above the species level is conceptual in the
mind of humans, and is pure semantics and does not exist in nature. There
are no subgenera, genera, subfamilies, families or orders scuttling around out
there on the ground, or questing on the vegetation or attached to hosts. But,
and this is where the crux of the matter lies, humans love organization, and
what better way to organize than to put objects similar in appearance in a
compartment with a particular name. However, when you convene a group of

The publication of this list has been facilitated by the International Consortium on Ticks
and Tick-borne Diseases (ICTTD-2) coordinated by Professor Frans Jongejan.
Author for correspondence (E-mail: ivan.horak@up.ac.za)
28

people to organize almost anything, there will never be complete agreement

on the correct arrangement of the entities being organized, and more espe-
cially so if instead of working in a group each person works in isolation. We
are well aware of differences in opinion, even amongst ourselves, but with
considerable discussion and some concessions we have put together a list of
tick names (Table 1) that we trust fastidious organizers will be able to live
with until someone arranges a better one.
In assembling this list we have had the advantage of having three previous
lists, containing the names of all known tick species at the various times
of their compilation, to consult (Neumann, 1911; Keirans, 1992; Camicas
et al., 1998), as well as a checklist of species published between 1973 and
1997 (Keirans and Robbins, 1999). We have used the systematic relation-
ships of the soft ticks proposed by Klompen and Oliver (1993) in compiling
the list of names for the Argasidae. These authors looked at the entire fam-
ily and the cladistic analysis of the 83 characters they used indicated that
there were only four genera, namely Argas, Carios, Ornithodoros and Oto-
bius within this family and the available evidence did not permit recognition
of subgenera in the latter three genera. This implies that the genera Alve-
onasus, Argas, Ogadenus, Persicargas, Proknekalia and Secretargas are now
subgenera of Argas while the genera Alectorobius, Antricola, Carios, Chirop-
terargas, Nothoaspis, Parantricola, Reticulinasus, and Subparmatus become
synonyms of Carios. One of us (JLC) does not fully agree with this classi-
fication of the Argasidae, and in particular with the status accorded to the
name Carios. However, we all agree that the systematic grouping of this
family requires further molecular taxonomic study before it can be settled
satisfactorily. The acceptance of the classification currently proposed presents
us with the problem of homonymy. The catalogue of Camicas et al. (1998)
lists Alveonasus cooleyi (McIvor, 1941) as well as Argas cooleyi Kohls and
Hoogstraal, 1960, and consequently when these genera are made subgenera
of Argas we now have two ticks named Argas cooleyi. This anomaly will
obviously have to be addressed some time in the future.
In the case of the Ixodidae we have incorporated the recommendations of
two recent publications. In one all the ticks within the genus Rhipicephalus
were examined and, but for two undescribed species, were assigned specific
names (Walker et al., 2000). In the other the classification of ticks of the genus
Aponomma has been rearranged and the more primitive members have been
transferred to the subgenus Bothriocroton Keirans, King and Sharrad, 1994,
which has been raised to generic rank, and the remainder to the genus
Amblyomma (Klompen et al., 2002). The genus Aponomma thus in part be-
comes a synonym of Bothriocroton and in part a synonym of Amblyomma.
An even more radical change that we have made is placing the five species
in the genus Boophilus in the genus Rhipicephalus. We do so with trepidation,
 29

but the current evidence in support of this change is overwhelming (Murrell

et al., 2000, 2001; Beati and Keirans, 2001), and in effect we are following
a proposal already made towards the end of the 19th century by Neumann
(1897). The genus Boophilus probably contains the globally best-known tick
of all, namely Boophilus microplus and arguably the best-known African
tick, Boophilus decoloratus. To avoid confusion, and even disbelief, amongst
fieldworkers concerning this name change, we have retained Boophilus as a
subgeneric epithet, and these ticks now become Rhipicephalus (Boophilus)
microplus and Rhipicephalus (Boophilus) decoloratus. In addition we have
not buried them in alphabetical sequence within the list of species belonging
to the genus Rhipicephalus, but have placed them separately at the head of
this list. This way, even if investigators continue to use Boophilus, everyone
will still know to what ticks they are referring.
We have added the names of four argasid ticks and five ixodid ticks, de-
scribed since 1996, to the list and that of one ixodid species omitted from
the previous list. We have not furnished sub-specific names in our list, except
where these are of importance in assisting with the identification of partic-
ular ticks. Even a cursory comparison of our list with the earlier lists will
reveal that we have again organized things differently. This may infuriate
zoologists, parasitologists and veterinarians who are not systematists, but
who work within the discipline of acarology and prefer classifications as
well as binominal epithets, and their spelling, to be cast in stone. For this
we apologize, but hope that you will nevertheless find the list useful.

Table 1. List of valid names for the Argasidae, Ixodidae and the Nuttalliellidae

Argasidae
Argasinae
Argas
A. abdussalami Hoogstraal and McCarthy, 1965
A. acinus (Whittick, 1938)
A. africolumbae Hoogstraal, Kaiser, Walker, Ledger, Converse and Rice, 1975
A. arboreus Kaiser, Hoogstraal and Kohls, 1964
A. assimilis Teng and Song, 1983
A. beijingensis Teng, 1983
A. beklemischevi Pospelova-Shtrom, Vasileva and Semashko, 1963
A. brevipes Banks, 1908
A. brumpti Neumann, 1907
A. bureschi Dryenski, 1957
A. canestrinii (Birula, 1895)
A. cooleyi (McIvor, 1941)
30
Table 1. (continued)

A. cooleyi Kohls and Hoogstraal, 1960

A. cucumerinus Neumann, 1901
A. dalei Clifford, Keirans, Hoogstraal and Corwin, 1976
A. delanoei (Roubaud and Colas-Belcour, 1931)
A. dulus Keirans, Clifford and Capriles, 1971
A. eboris (Theiler, 1959)
A. echinops Hoogstraal, Uilenberg and Blanc, 1967
A. falco Kaiser and Hoogstraal, 1974
A. foleyi (Parrot, 1928)
A. giganteus Kohls and Clifford, 1968
A. gilcolladoi Estrada-Pea, Lucientes and Sanchez, 1987
A. hermanni Audouin, 1827
A. himalayensis Hoogstraal and Kaiser, 1973
A. hoogstraali Morel and Vassiliades, 1965
A. japonicus Yamaguti, Clifford and Tipton, 1968
A. lagenoplastis Froggatt, 1906
A. lahorensis (Neumann, 1908)
A. latus Filippova, 1961
A. lowryae Kaiser and Hoogstraal, 1975
A. macrostigmatus Filippova, 1961
A. magnus Neumann, 1896
A. miniatus Koch, 1844
A. monachus Keirans, Radovsky and Clifford, 1973
A. monolakensis Schwan, Corwin and Brown, 1992
A. moreli Keirans, Hoogstraal and Clifford, 1979
A. neghmei Kohls and Hoogstraal, 1961
A. nullarborensis Hoogstraal and Kaiser, 1973
A. peringueyi (Bedford and Hewitt, 1925)
A. persicus (Oken, 1818)
A. peusi (Schulze, 1943)
A. polonicus Siuda, Hoogstraal, Clifford and Wassef, 1979
A. radiatus Railliet, 1893
A. reflexus (Fabricius, 1794)
A. ricei Hoogstraal, Kaiser, Clifford and Keirans, 1975
A. robertsi Hoogstraal, Kaiser and Kohls, 1968
A. sanchezi Dug`es, 1887
A. streptopelia Kaiser, Hoogstraal and Horner, 1970
A. striatus Bedford, 1932
A. theilerae Hoogstraal and Kaiser, 1970
 31
Table 1. (continued)

A. transgariepinus White, 1846

A. tridentatus Filippova, 1961
A. vansomereni (Keirans, Hoogstraal and Clifford, 1977)
A. vulgaris Filippova, 1961 (?=A. delicatus Neumann, 1910)
A. walkerae Kaiser and Hoogstraal, 1969
A. zumpti Hoogstraal, Kaiser and Kohls, 1968

Ornithodorinae
Carios
C. amblus (Chamberlin, 1920)
C. aragaoi (Fonseca, 1960)
C. armasi (de la Cruz and Estrada-Pea, 1995)
C. australiensis (Kohls and Hoogstraal, 1962)
C. azteci (Matheson, 1935)
C. batuensis (Hirst, 1929)
C. boueti (Roubaud and Colas-Belcour, 1933)
C. brodyi (Matheson, 1935)
C. camicasi (Sylla, Cornet and Marchand, 1997)
C. capensis (Neumann, 1901)
C. casebeeri (Jones and Clifford, 1972)
C. centralis (de la Cruz and Estrada-Pea, 1995)
C. cernyi (de la Cruz, 1978)
C. ceylonensis (Hoogstraal and Kaiser, 1968)
C. cheikhi (Vermeil, Marjolet and Vermeil, 1997)
C. chironectes (Jones and Clifford, 1972)
C. chiropterphila (Dhanda and Rajagopalan, 1971)
C. clarki (Jones and Clifford, 1972)
C. collocaliae (Hoogstraal, Kadarsan, Kaiser and Van Peenen, 1974)
C. concanensis (Cooley and Kohls, 1941)
C. confusus (Hoogstraal, 1955)
C. coniceps (Canestrini, 1890)
C. coprophilus (McIntosh, 1935)
C. cordiformis (Hoogstraal and Kohls, 1967)
C. cyclurae (de la Cruz, 1984)
C. darwini (Kohls, Clifford and Hoogstraal, 1969)
C. daviesi (Kaiser and Hoogstraal, 1973)
C. denmarki (Kohls, Sonenshine and Clifford, 1965)
C. dewae (Kaiser and Hoogstraal, 1974)
y, 1967)
C. dusbabeki (Cern
32
Table 1. (continued)

C. dyeri (Cooley and Kohls, 1940)

C. echimys (Kohls, Clifford and Jones, 1969)
C. elongatus (Kohls, Sonenshine and Clifford, 1965)
C. eptesicus (Kohls, Clifford and Jones, 1969)
C. faini (Hoogstraal, 1960)
C. galapagensis (Kohls, Clifford and Hoogstraal, 1969)
C. granasi (de la Cruz, 1973)
C. habanensis (de la Cruz, 1976)
C. hadiae Klompen, Keirans and Durden, 1995
C. hasei (Schulze, 1935)
C. hummelincki (de la Cruz and Estrada-Pea, 1995)
C. jerseyi Klompen and Grimaldi, 2001
C. jul (Schulze, 1940)
C. kelleyi (Cooley and Kohls, 1941)
C. kohlsi (Guglielmone and Keirans, 2002)
C. macrodermae (Hoogstraal, Moorhouse, Wolf and Wassef, 1977)
C. madagascariensis (Hoogstraal, 1962)
C. marginatus (Banks, 1910)
C. marinkellei (Kohls, Clifford and Jones, 1969)
C. maritimus (Vermeil and Marguet, 1967)
C. marmosae (Jones and Clifford, 1972)
C. martelorum (de la Cruz, 1978)
C. mexicanus (Hoffmann, 1959)
C. mimon (Kohls, Clifford and Jones, 1969)
C. mormoops (Kohls, Clifford and Jones, 1969)
C. muesebecki (Hoogstraal, 1969)
C. multisetosus Klompen, Keirans and Durden, 1995
C. naomiae (de la Cruz, 1978)
y and Dusbabek, 1967)
C. natalinus (Cern
C. occidentalis (de la Cruz, 1978)
C. papuensis Klompen, Keirans and Durden, 1995
C. peropteryx (Kohls, Clifford and Jones, 1969)
C. peruvianus (Kohls, Clifford and Jones, 1969)
C. piriformis (Warburton, 1918)
C. puertoricensis (Fox, 1947)
C. pusillus (Kohls, 1950)
C. reddelli (Keirans and Clifford, 1975)
C. rennellensis (Clifford and Sonenshine, 1962)
C. rossi (Kohls, Sonenshine and Clifford, 1965)
 33
Table 1. (continued)

C. rudis (Karsch, 1880)

C. salahi (Hoogstraal, 1953)
C. sawaii (Kitaoka and Suzuki, 1973)
C. setosus (Kohls, Clifford and Jones, 1969)
C. siboneyi (de la Cruz and Estrada-Pea, 1995)
y, 1967)
C. silvai (Cern
C. sinensis (Jeu and Zhu, 1982)
C. solomonis (Dumbleton, 1959)
C. spheniscus (Hoogstraal, Wassef, Hays and Keirans, 1985)
C. stageri (Cooley and Kohls, 1941)
y and Dusbabek, 1967)
C. tadaridae (Cern
C. talaje (Guerin-Meneville, 1849)
C. tiptoni (Jones and Clifford, 1972)
C. tuttlei (Jones and Clifford, 1972)
C. vespertilionis Latreille, 1796
C. viguerasi (Cooley and Kohls, 1941)
C. yumatensis (Cooley and Kohls, 1941)
C. yunkeri (Keirans, Clifford and Hoogstraal, 1984)
Ornithodoros
O. alactagalis Issaakjan, 1936
O. antiquus Poinar, 1995 (a fossil species)
O. apertus Walton, 1962
O. arenicolous Hoogstraal, 1953
O. asperus Warburton, 1918
O. brasiliensis Arago, 1923
O. cholodkovskyi Pavlovsky, 1930
O. compactus Walton, 1962
O. coriaceus Koch, 1844
O. eremicus Cooley and Kohls, 1941
O. erraticus (Lucas, 1849)
O. furcosus Neumann, 1908
O. graingeri Heisch and Guggisberg, 1953
O. grenieri Klein, 1965
O. gurneyi Warburton, 1926
O. hermsi Wheeler, Herms and Meyer, 1935
O. indica Rau and Rao, 1971
O. knoxjonesi Jones and Clifford, 1972
O. macmillani (Hoogstraal and Kohls, 1966)
O. marocanus Velu, 1919
34
Table 1. (continued)

O. moubata (Murray, 1877)

O. nattereri Warburton, 1927
O. nicollei Mooser, 1932
O. normandi Larrousse, 1923
O. parkeri Cooley, 1936
O. porcinus avivora Walton, 1979
O. porcinus domesticus Walton, 1962
O. porcinus porcinus Walton,1962
O. procaviae Theodor and Costa, 1960
O. rostratus Arago, 1911
O. savignyi (Audouin, 1827)
O. sonrai Sautet and Witkowski, 1943
O. steini (Schulze, 1935)
O. tartakovskyi Olenev, 1931
O. tholozani Laboulb`ene and Megnin, 1882
O. transversus (Banks, 1902)
O. turicata (Dug`es, 1876)
O. zumpti Heisch and Guggisberg, 1953
Otobius
O. lagophilus Cooley and Kohls, 1940
O. megnini (Dug`es, 1883)
O. sparnus (Kohls and Clifford, 1963)

Ixodidae
Ixodinae
Ixodes
I. abrocomae Lahille, 1917
I. acuminatus Neumann, 1901
I. acutitarsus (Karsch, 1880)
I. affinis Neumann, 1899
I. albignaci Uilenberg and Hoogsraal, 1969
I. alluaudi Neumann, 1913
I. amarali Fonseca, 1935
I. amersoni Kohls, 1966
I. anatis Chilton, 1904
I. andinus Kohls, 1956
I. angustus Neumann, 1899
I. antechini Roberts, 1960
I. apronophorus Schulze, 1924
 35
Table 1. (continued)

I. arabukiensis Arthur, 1959

I. aragaoi Fonseca, 1935
I. arboricola Schulze and Schlottke, 1930
I. arebiensis Arthur, 1956
I. asanumai Kitaoka, 1973
I. aulacodi Arthur, 1956
I. auriculaelongae Arthur, 1958
I. auritulus Neumann, 1904 (=I. auritulus zealandicus Dumbleton, 1953)
I. australiensis Neumann, 1904
I. baergi Cooley and Kohls, 1942
I. bakeri Arthur and Clifford, 1961
I. banksi Bishopp, 1911
I. bedfordi Arthur, 1959
I. bequaerti Cooley and Kohls, 1945
I. berlesei Birula, 1895
I. bivari Santos Dias, 1990
I. boliviensis Neumann, 1904
I. brewsterae Keirans, Clifford and Walker, 1982
I. browningi Arthur, 1956
I. brumpti Morel, 1965
I. brunneus Koch, 1844
I. calcarhebes Arthur and Zulu, 1980
I. caledonicus Nuttall, 1910
I. canisuga Johnston, 1849
y, 1966
I. capromydis Cern
I. catherinei Keirans, Clifford and Walker, 1982
I. cavipalpus Nuttall and Warburton, 1908
I. ceylonensis Kohls, 1950
I. chilensis Kohls, 1956
I. colasbelcouri Arthur, 1957
I. collocaliae Schulze, 1937
I. columnae Takada and Fujita, 1992
I. conepati Cooley and Kohls, 1943
I. confusus Roberts, 1960
I. cookei Packard, 1869
I. cooleyi Arago and Fonseca, 1951
I. copei Wilson, 1980
I. cordifer Neumann, 1908
36
Table 1. (continued)

I. cornuae Arthur, 1960

I. cornuatus Roberts, 1960
I. corwini Keirans, Clifford and Walker, 1982
I. crenulatus Koch, 1844
I. cuernavacensis Kohls and Clifford, 1966
I. cumulatimpunctatus Schulze, 1943
I. dampfi Cooley, 1943
I. daveyi Nuttall, 1913
I. dawesi Arthur, 1956
I. dendrolagi Wilson, 1967
I. dentatus Marx, 1899
I. diomedeae Arthur, 1958
I. diversifossus Neumann, 1899
I. djaronensis Neumann, 1907
I. domerguei Uilenberg and Hoogstraal, 1965
I. downsi Kohls, 1957
I. drakensbergensis Clifford, Theiler and Baker, 1975
I. eadsi Kohls and Clifford, 1964
I. eastoni Keirans and Clifford, 1983
I. eichhorni Nuttall, 1916
I. eldaricus Dzhaparidze, 1950
I. elongatus Bedford, 1929
I. eudyptidis Maskell, 1885
I. euplecti Arthur, 1958
I. evansi Arthur, 1956
I. fecialis Warburton and Nuttall, 1909
I. festai Rondelli, 1926
y, 1961
I. filippovae Cern
I. fossulatus Neumann, 1899
I. frontalis Panzer, 1798
I. fuscipes Koch, 1844
I. galapagoensis Clifford and Hoogstraal, 1980
I. ghilarovi Filippova and Panova, 1988
I. gibbosus Nuttall, 1916
I. granulatus Supino, 1897
I. gregsoni Lindquist, Wu and Redner, 1999
I. guatemalensis Kohls, 1956
I. hearlei Gregson, 1941
 37
Table 1. (continued)

I. heinrichi Arthur, 1962

I. hexagonus Leach, 1815
I. himalayensis Dhanda and Kulkarni, 1969
I. hirsti Hassall, 1931
I. holocyclus Neumann, 1899
I. hoogstraali Arthur, 1955
I. howelli Cooley and Kohls, 1938
I. hyatti Clifford, Hoogstraal and Kohls, 1971
I. hydromyidis Swan, 1931
I. jacksoni Hoogstraal, 1967
I. jellisoni Cooley and Kohls, 1938
I. jonesae Kohls, Sonenshine and Clifford, 1969
I. kaiseri Arthur, 1957
I. kashimiricus Pomerantsev, 1948
I. kazakstani Olenev and Sorokoumov, 1934
I. kerguelenensis Andre and Colas-Belcour, 1942
I. kingi Bishopp, 1911
I. kohlsi Arthur, 1955
I. kopsteini Oudemans, 1926
I. kuntzi Hoogstraal and Kohls, 1965
I. laguri Olenev, 1929 sensu Olenev, 1931
I. lasallei Mendez Arocha and Ortiz, 1958
I. latus Arthur, 1958
I. laysanensis Wilson, 1964
I. lemuris Arthur, 1958
I. lewisi Arthur, 1965
I. lividus Koch, 1844
I. longiscutatus Boero, 1944
I. loricatus Neumann, 1899
I. loveridgei Arthur, 1958
I. luciae Senevet, 1940
I. lunatus Neumann, 1907
I. luxuriosus Schulze, 1932
I. macfarlanei Keirans, Clifford and Walker, 1982
I. malayensis Kohls, 1962
I. marmotae Cooley and Kohls, 1938
I. marxi Banks, 1908
I. maslovi Emelyanova and Kozlovskaya, 1967
I. matopi Spickett, Keirans, Norval and Clifford, 1981
38
Table 1. (continued)

I. mexicanus Cooley and Kohls, 1942

I. minor Neumann, 1902
I. minutae Arthur, 1959
I. mitchelli Kohls, Clifford and Hoogstraal, 1970
I. monospinosus Saito, 1968
I. montoyanus Cooley, 1944
I. moreli Arthur, 1957
I. moscharius Teng, 1982
I. moschiferi Nemenz, 1968
I. muniensis Arthur and Burrow, 1957
I. muris Bishopp and Smith, 1937
I. murreleti Cooley and Kohls, 1945
I. myospalacis Teng, 1986
I. myotomys Clifford and Hoogstraal, 1970
I. myrmecobii Roberts, 1962
I. nairobiensis Nuttall, 1916
I. nchisiensis Arthur, 1958
I. nectomys Kohls, 1956
I. neitzi Clifford, Walker and Keirans, 1977
I. nesomys Uilenberg and Hoogstraal, 1969
I. neuquenensis Ringuelet, 1947
I. nicolasi Santos Dias, 1982
I. nipponensis Kitaoka and Saito, 1967
I. nitens Neumann, 1904
I. nuttalli Lahille, 1913
I. nuttallianus Schulze, 1930
I. occultus Pomerantsev, 1946
I. ochotonae Gregson, 1941
I. okapiae Arthur, 1956
I. oldi Nuttall, 1913
I. ornithorhynchi Lucas, 1846
I. ovatus Neumann, 1899
I. pacificus Cooley and Kohls, 1943
I. pararicinus Keirans and Clifford, 1985
I. pavlovskyi Pomerantsev, 1946
I. percavatus Neumann, 1906
I. peromysci Augustson, 1940
I. persulcatus Schulze, 1930
 39
Table 1. (continued)

I. petauristae Warburton, 1933

I. philipi Keirans and Kohls, 1970
I. pilosus Koch, 1844
I. pomerantzi Kohls, 1956
I. pomeranzevi Serdyukova, 1941
I. priscicollaris Schulze, 1932
I. procaviae Arthur and Burrow, 1957
I. prokopjevi (Emelyanova, 1979)
I. radfordi Kohls, 1948
I. rageaui Arthur, 1958
I. randrianasoloi Uilenberg and Hoogstraal, 1969
I. rangtangensis Teng, 1973
I. rasus Neumann, 1899
I. redikorzevi Olenev, 1927
I. rhabdomysae Arthur, 1959
I. ricinus (Linnaeus, 1758)
I. rothschildi Nuttall and Warburton, 1911
I. rotundatus Arthur, 1958
I. rubicundus Neumann, 1904
I. rubidus Neumann, 1901
I. rugicollis Schulze and Schlottke, 1930
I. rugosus Bishopp, 1911
I. sachalinensis Filippova, 1971
I. scapularis Say, 1821
I. schillingsi Neumann, 1901
I. schulzei Arago and Fonseca, 1951
I. sculptus Neumann, 1904
I. semenovi Olenev, 1929
I. shahi Clifford, Hoogstraal and Kohls, 1971
I. siamensis Kitaoka and Suzuki, 1983
I. sigelos Keirans, Clifford and Corwin, 1976
I. signatus Birula, 1895
I. simplex Neumann, 1906
I. sinaloa Kohls and Clifford, 1966
I. sinensis Teng, 1977
I. soricis Gregson, 1942
I. spinae Arthur, 1958
I. spinicoxalis Neumann, 1899
I. spinipalpis Hadwen and Nuttall, 1916
40
Table 1. (continued)

I. steini Schulze, 1932

I. stilesi Neumann, 1911
I. stromi Filippova, 1957
I. subterranus Filippova, 1961
I. taglei Kohls, 1969
I. tamaulipas Kohls and Clifford, 1966
I. tancitarius Cooley and Kohls, 1942
I. tanuki Saito, 1964
I. tapirus Kohls, 1956
I. tasmani Neumann, 1899
I. tecpanensis Kohls, 1956
I. texanus Banks, 1909
I. theilerae Arthur, 1953
I. thomasae Arthur and Burrow, 1957
I. tiptoni Kohls and Clifford, 1962
I. tovari Cooley, 1945
I. transvaalensis Clifford and Hoogstraal, 1966
I. trianguliceps Birula, 1895
I. trichosuri Roberts, 1960
I. tropicalis Kohls, 1956
I. turdus Nakatsuji, 1942
I. ugandanus Neumann, 1906
I. unicavatus Neumann, 1908
I. uriae White, 1852
I. vanidicus Schulze, 1943
I. venezuelensis Kohls, 1953
I. ventalloi Gil Collado, 1936
I. vespertilionis Koch, 1844
I. vestitus Neumann, 1908
I. victoriensis Nuttall, 1916
I. walkerae Clifford, Kohls and Hoogstraal, 1968
I. werneri Kohls, 1950
I. woodi Bishopp, 1911
I. zaglossi Kohls, 1960
I. zairensis Keirans, Clifford and Walker, 1982
I. zumpti Arthur, 1960

Amblyomminae
Amblyomma
A. albolimbatum Neumann, 1907
A. albopictum Neumann, 1899
 41
Table 1. (continued)

A. americanum (Linnaeus, 1758)

A. antillorum Kohls, 1969
A. argentinae Neumann, 1905
A. astrion Dnitz, 1909
A. aureolatum (Pallas, 1772)
A. auricularium (Conil, 1878)
A. australiense Neumann, 1905
A. babirussae Schulze, 1933
A. boulengeri Hirst and Hirst, 1910
A. brasiliense Arago, 1908
A. breviscutatum Neumann, 1899
A. cajennense (Fabricius, 1787)
A. calabyi Roberts, 1963
A. calcaratum Neumann, 1899
A. chabaudi Rageau, 1964
A. clypeolatum Neumann, 1899
A. coelebs Neumann, 1899
A. cohaerens Dnitz, 1909
A. compressum (Macalister, 1872)
A. cordiferum Neumann, 1899
A. crassum Robinson, 1926
A. crenatum Neumann, 1899
A. cruciferum Neumann, 1901
A. darwini Hirst and Hirst, 1910
A. dissimile Koch, 1844
A. dubitatum Neumann, 1899
A. eburneum Gerstcker, 1873
A. echidnae Roberts, 1953
A. extraoculatum Neumann, 1899
A. falsomarmoreum Tonelli-Rondelli, 1935
A. fulvum Neumann, 1899
A. geayi Neumann, 1899
A. gemma Dnitz, 1909
A. geochelone Durden, Keirans and Smith, 2002
A. geoemydae (Cantor, 1847)
A. glauerti Keirans, King and Sharrad, 1994
A. goeldii Neumann, 1899
A. hainanense Teng, 1981
A. hebraeum Koch, 1844
A. helvolum Koch, 1844
A. hirtum Neumann, 1906
42
Table 1. (continued)

A. humerale Koch, 1844

A. imitator Kohls, 1958
A. incisum Neumann, 1906
A. inornatum (Banks, 1909)
A. integrum Karsch, 1879
A. javanense (Supino, 1897)
A. laticaudae Warburton, 1933
A. lepidum Dnitz, 1909
A. limbatum Neumann, 1899
A. loculosum Neumann, 1907
A. longirostre (Koch, 1844)
A. macfarlandi Keirans, Hoogstraal and Clifford, 1973
A. macropi Roberts, 1953
A. maculatum Koch, 1844
A. marmoreum Koch, 1844
A. moreliae (Koch, 1867)
A. moyi Roberts, 1953
A. multipunctum Neumann, 1899
A. naponense (Packard, 1869)
A. neumanni Ribaga, 1902
A. nitidum Hirst and Hirst, 1910
A. nodosum Neumann, 1899
A. nuttalli Dnitz, 1909
A. oblongoguttatum Koch, 1844
A. ovale Koch, 1844
A. pacae Arago, 1911
A. papuanum Hirst, 1914
A. parvitarsum Neumann, 1901
A. parvum Arago, 1908
A. paulopunctatum Neumann, 1899
A. pecarium Dunn, 1933
A. personatum Neumann, 1901
A. pictum Neumann, 1906
A. pilosum Neumann, 1899
A. pomposum Dnitz, 1909
A. postoculatum Neumann, 1899
A. pseudoconcolor Arago, 1908
A. pseudoparvum Guglielmone, Mangold and Keirans, 1990
A. quadricavum (Schulze, 1941)
 43
Table 1. (continued)

A. rhinocerotis (de Geer, 1778)

A. robinsoni Warburton, 1927
A. rotundatum Koch, 1844
A. sabanerae Stoll, 1890
A. scalpturatum Neumann, 1906
A. scutatum Neumann, 1899
A. sparsum Neumann, 1899
A. splendidum Giebel, 1877
A. squamosum Kohls, 1953
A. supinoi Neumann, 1905
A. sylvaticum (de Geer, 1778)
A. tapirellum Dunn, 1933
A. testudinarium Koch, 1844
A. tholloni Neumann, 1899
A. tigrinum Koch, 1844
A. torrei Perez Vigueras, 1934
A. triguttatum ornatissimum Roberts, 1962
A. triguttatum queenslandense Roberts, 1962
A. triguttatum rosei Roberts, 1962
A. triguttatum triguttatum Koch, 1844
A. triste Koch, 1844
A. tuberculatum Marx, 1894
A. usingeri Keirans, Hoogstraal and Clifford, 1973
A. variegatum (Fabricius, 1794)
A. varium Koch, 1844
A. vikirri Keirans, Bull and Duffield, 1996
A. williamsi Banks, 1924
Amblyomma (Former Aponomma, now synonyms of Amblyomma)
A. arcanum (Karsch, 1879)
A. crassipes (Neumann, 1901)
A. elaphense (Price, 1959)
A. exornatum Koch, 1844
A. fimbriatum Koch, 1844
A. flavomaculatum (Lucas, 1846)
A. fuscolineatum (Lucas, 1847)
A. gervaisi (Lucas, 1847)
A. inopinatum (Santos Dias, 1989)
A. komodoense (Oudemans, 1928)
A. kraneveldi (Anastos, 1956)
44

Table 1. (continued)

A. latum Koch, 1844

A. orlovi (Kolonin, 1992)
A. oudemansi (Neumann, 1910)
A. pattoni (Neumann, 1910)
A. soembawense (Anastos, 1956)
A. sphenodonti (Dumbleton, 1943)
A. transversale (Lucas, 1844)
A. trimaculatum (Lucas, 1878)
A. varanense (Supino, 1897)

Bothriocrotoninae
Bothriocroton
B. auruginans (Schulze, 1936)
B. concolor (Neumann, 1899)
B. glebopalma (Keirans, King and Sharrad, 1994)
B. hydrosauri (Denny, 1843)
B. undatum (Fabricius, 1775)

Haemaphysalinae
Haemaphysalis
H. aborensis Warburton, 1913
H. aciculifer Warburton, 1913
H. aculeata Lavarra, 1904
H. adleri Feldman-Muhsam, 1951
H. anomala Warburton, 1913
H. anomaloceraea Teng and Cui, 1984
H. anoplos Hoogstraal, Uilenberg and Klein, 1967
H. aponommoides Warburton, 1913
H. asiatica (Supino, 1897)
H. atheruri Hoogstraal, Trapido and Kohls, 1965
H. bancrofti Nuttall and Warburton, 1915
H. bandicota Hoogstraal and Kohls, 1965
H. bartelsi Schulze, 1938
H. bequaerti Hoogstraal, 1956
H. birmaniae Supino, 1897
H. bispinosa Neumann, 1897
H. borneata Hoogstraal, 1971
H. bremneri Roberts, 1963
 45
Table 1. (continued)

H. calcarata Neumann, 1902

H. calva Nuttall and Warburton, 1915
H. campanulata Warburton, 1908
H. canestrinii (Supino, 1897)
H. capricornis Hoogstraal, 1966
H. caucasica Olenev, 1928
H. celebensis Hoogstraal, Trapido and Kohls, 1965
H. chordeilis (Packard, 1869)
H. colasbelcouri (Santos Dias, 1958)
H. concinna Koch, 1844
H. cooleyi Bedford, 1929
H. cornigera Neumann, 1897
H. cornupunctata Hoogstraal and Varma, 1962
H. cuspidata Warburton, 1910
H. dangi Phan Trong, 1977
y and Hoogstraal, 1977
H. danieli Cern
H. darjeeling Hoogstraal and Dhanda, 1970
H. davisi Hoogstraal, Dhanda and Bhat, 1970
H. demidovae Emelyanova, 1978
H. doenitzi Warburton and Nuttall, 1909
H. elliptica (Koch, 1844)
H. elongata Neumann, 1897
H. erinacei erinacei Pavesi, 1884
H. erinacei ornata Feldman-Muhsam, 1956
H. erinacei taurica Pospelova-Shtrom, 1940
H. erinacei turanica Pospelova-Shtrom, 1940
H. eupleres Hoogstraal, Kohls and Trapido, 1965
H. filippovae Bolotin, 1979
H. flava Neumann, 1897
H. formosensis Neumann, 1913
H. fossae Hoogstraal, 1953
H. fujisana Kitaoka, 1970
H. garhwalensis Dhanda and Bhat, 1968
H. goral Hoogstraal, 1970
H. grochovskajae Kolonin, 1992
H. heinrichi Schulze, 1939
H. hirsuta Hoogstraal, Trapido and Kohls, 1966
H. hispanica Gil Collado, 1938
46
Table 1. (continued)

H. hoodi Warburton and Nuttall, 1909

H. hoogstraali Kohls, 1950
H. houyi Nuttall and Warburton, 1915
H. howletti Warburton, 1913
H. humerosa Warburton and Nuttall, 1909
H. hylobatis Schulze, 1933
H. hyracophila Hoogstraal, Walker and Neitz, 1971
H. hystricis Supino, 1897
H. ias Nakamura and Yajima, 1937
H. indica Warburton, 1910
H. indoflava Dhanda and Bhat, 1968
H. inermis Birula, 1895
H. intermedia Warburton and Nuttall, 1909
H. japonica douglasi Nuttall and Warburton, 1915
H. japonica japonica Warburton, 1908
H. juxtakochi Cooley, 1946
H. kadarsani Hoogstraal and Wassef, 1977
H. kashmirensis Hoogstraal and Varma, 1962
H. kinneari Warburton, 1913
H. kitaokai Hoogstraal, 1969
H. koningsbergeri Warburton and Nuttall, 1909
H. kopetdaghica Kerbabaev, 1962
H. kutchensis Hoogstraal and Trapido, 1963
H. kyasanurensis Trapido, Hoogstraal and Rajagopalan, 1964
H. lagostrophi Roberts, 1963
H. lagrangei Larrousse, 1925
H. laocayensis Phan Trong, 1977
H. leachi (Audouin, 1826)
H. lemuris Hoogstraal, 1953
H. leporispalustris (Packard, 1869)
H. lobachovi Kolonin, 1995
H. longicornis Neumann, 1901
H. luzonensis Hoogstraal and Parrish, 1968
H. madagascariensis Colas-Belcour and Millot, 1948
H. mageshimaensis Saito and Hoogstraal, 1973
H. megalaimae Rajagopalan, 1963
H. megaspinosa Saito, 1969
H. menglaensis Pang, Chen and Xiang, 1982
H. minuta Kohls, 1950
 47
Table 1. (continued)

H. mjoebergi Warburton, 1926

H. montgomeryi Nuttall, 1912
H. moreli Camicas, Hoogstraal and El Kammah, 1972
H. moschisuga Teng, 1980
H. muhsamae Santos Dias, 1954
H. nadchatrami Hoogstraal, Trapido and Kohls, 1965
H. nepalensis Hoogstraal, 1962
H. nesomys Hoogstraal, Uilenberg and Klein, 1966
H. norvali Hoogstraal and Wassef, 1983
H. novaeguineae Hirst, 1914
H. obesa Larrousse, 1925
H. obtusa Dnitz, 1910
H. orientalis Nuttall and Warburton, 1915
H. ornithophila Hoogstraal and Kohls, 1959
H. palawanensis Kohls, 1950
H. papuana Thorell, 1883
H. paraleachi Camicas, Hoogstraal and El Kammah, 1983
H. paraturturis Hoogstraal, Trapido and Rebello, 1963
H. parmata Neumann, 1905
H. parva Neumann, 1897
H. pavlovskyi Pospelova-Shtrom, 1935
H. pedetes Hoogstraal, 1972
H. pentalagi Pospelova-Shtrom, 1935
H. petrogalis Roberts, 1970
H. phasiana Saito, Hoogstraal and Wassef, 1974
H. pospelovashtromae Hoogstraal, 1966
H. primitiva Teng, 1982
H. psalistos Hoogstraal, Kohls and Parrish, 1967
H. punctaleachi Camicas, Hoogstraal and El Kammah, 1973
H. punctata Canestrini and Fanzago, 1878
H. qinghaiensis Teng, 1980
H. quadriaculeata Kolonin, 1992
H. ramachandrai Dhanda, Hoogstraal and Bhat, 1970
H. ratti Kohls, 1948
H. renschi Schulze, 1933
H. roubaudi Toumanoff, 1940
H. rugosa Santos Dias, 1956
H. rusae Kohls, 1950
H. sambar Hoogstraal, 1971
48
Table 1. (continued)

H. sciuri Kohls, 1950

H. semermis Neumann, 1901
H. shimoga Trapido and Hoogstraal, 1964
H. silacea Robinson, 1912
H. silvafelis Hoogstraal and Trapido, 1963
H. simplex Neumann, 1897
H. simplicima Hoogstraal and Wassef, 1979
H. sinensis Zhang, 1981
H. spinigera Neumann, 1897
H. spinulosa Neumann, 1906
H. subelongata Hoogstraal, 1953
H. subterra Hoogstraal, El Kammah and Camicas, 1992
H. sulcata Canestrini and Fanzago, 1878
H. sumatraensis Hoogstraal, El Kammah, Kadarsan and Anastos, 1971
H. sundrai Sharif, 1928
H. suntzovi Kolonin, 1993
H. susphilippensis Hoogstraal, Kohls and Parrish, 1968
H. taiwana Sugimoto, 1936
H. tauffliebi Morel, 1965
H. theilerae Hoogstraal, 1953
H. tibetensis Hoogstraal, 1965
H. tiptoni Hoogstraal, 1953
H. toxopei Warburton, 1927
H. traguli Oudemans, 1928
H. traubi Kohls, 1955
H. turturis Nuttall and Warburton, 1915
H. verticalis Itagaki, Noda and Yamaguchi, 1944
H. vidua Warburton and Nuttall, 1909
H. vietnamensis Hoogstraal and Wilson, 1966
H. warburtoni Nuttall, 1912
H. wellingtoni Nuttall and Warburton, 1908
H. xinjiangensis Teng, 1980
H. yeni Toumanoff, 1944
H. zumpti Hoogstraal and El Kammah, 1974

Hyalomminae
Hyalomma
H. aegyptium (Linnaeus, 1758)
H. albiparmatum Schulze, 1919
H. anatolicum anatolicum Koch, 1844
 49
Table 1. (continued)

H. anatolicum excavatum Koch, 1844

H. arabica Pegram, Hoogstraal and Wassef, 1982
H. asiaticum asiaticum Schulze and Schlottke, 1930
H. asiaticum caucasicum Pomerantsev, Matikashvili and Lototsky, 1940
H. asiaticum kozlovi Olenev, 1931
H. brevipunctata Sharif, 1928
H. detritum detritum Schulze, 1919
H. detritum scupense Schulze, 1918
H. dromedarii Koch, 1844
H. erythraeum Tonelli-Rondelli, 1932
H. franchinii Tonelli-Rondelli, 1932
H. hussaini Sharif, 1928
H. hystricis Dhanda and Raja, 1974
H. impeltatum Schulze and Schlottke, 1930
H. impressum Koch, 1844
H. kumari Sharif, 1928
H. lusitanicum Koch, 1844
H. marginatum isaaci Sharif, 1928
H. marginatum marginatum Koch, 1844
H. marginatum rufipes Koch, 1844
H. nitidum Schulze, 1919
H. punt Hoogstraal, Kaiser and Pedersen, 1969
H. rhipicephaloides Neumann, 1901
H. schulzei Olenev, 1931
H. sinaii Feldman-Muhsam, 1960
H. truncatum Koch, 1844
H. turanicum Pomerantsev, 1946
Nosomma
N. monstrosum (Nuttall and Warburton, 1908)

Rhipicephalinae
Anomalohimalaya
A. cricetuli Teng and Huang, 1981
A. lamai Hoogstraal, Kaiser and Mitchell, 1970
A. lotozkyi Filippova and Panova, 1978
Cosmiomma
C. hippopotamensis (Denny, 1843)
Dermacentor
D. abaensis Teng, 1963
D. albipictus (Packard, 1869)
50
Table 1. (continued)

D. andersoni Stiles, 1908

D. asper Arthur, 1960
D. atrosignatus Neumann, 1906
D. auratus Supino, 1897
D. circumguttatus Neumann, 1897
D. compactus Neumann, 1901
D. dispar Cooley, 1937
D. dissimilis Cooley, 1947
D. everestianus Hirst, 1926
D. halli McIntosh, 1931
D. hunteri Bishopp, 1912
D. imitans Warburton, 1933
D. latus Cooley, 1937
D. marginatus (Sulzer, 1776)
D. montanus Filippova and Panova, 1974
D. nitens Neumann, 1897
D. niveus Neumann, 1897
D. nuttalli Olenev, 1928
D. occidentalis Marx, 1892
D. parumapertus Neumann, 1901
D. pavlovskyi Olenev, 1927
D. pomerantzevi Serdyukova, 1951
D. raskemensis Pomerantsev, 1946
D. reticulatus (Fabricius, 1794)
D. rhinocerinus (Denny, 1843)
D. silvarum Olenev, 1931
D. sinicus Schulze, 1932
D. steini Schulze, 1933
D. taiwanensis Sugimoto, 1935
D. ushakovae Filippova and Panova, 1987
D. variabilis (Say, 1821)
Margaropus
M. reidi Hoogstraal, 1956
M. wileyi Walker and Laurence, 1973
M. winthemi Karsch, 1879
Rhipicentor
R. bicornis Nuttall and Warburton, 1908
R. nuttalli Cooper and Robinson, 1908
 51
Table 1. (continued)

Rhipicephalus
R. (Boophilus) annulatus (Say, 1821)
R. (Boophilus) decoloratus (Koch, 1844)
R. (Boophilus) geigyi (Aeschlimann and Morel, 1965)
R. (Boophilus) kohlsi (Hoogstraal and Kaiser, 1960)
R. (Boophilus) microplus (Canestrini, 1888)

R. appendiculatus Neumann, 1901

R. aquatilis Walker, Keirans and Pegram, 1993
R. armatus Pocock, 1900
R. arnoldi Theiler and Zumpt, 1949
R. bequaerti Zumpt, 1949
R. bergeoni Morel and Balis, 1976
R. boueti Morel, 1957
R. bursa Canestrini and Fanzago, 1878
R. camicasi Morel, Mouchet and Rodhain, 1976
R. capensis Koch, 1844
R. carnivoralis Walker, 1966
R. complanatus Neumann, 1911
R. compositus Neumann, 1897
R. cuspidatus Neumann, 1906
R. deltoideus Neumann, 1910
R. distinctus Bedford, 1932
R. duttoni Neumann, 1907
R. dux Dnitz, 1910
R. evertsi evertsi Neumann, 1897
R. evertsi mimeticus Dnitz, 1910
R. exophthalmos Keirans and Walker, 1993
R. follis Dnitz, 1910
R. fulvus Neumann, 1913
R. gertrudae Feldman-Muhsam, 1960
R. glabroscutatum Du Toit, 1941
R. guilhoni Morel and Vassiliades, 1963
R. haemaphysaloides (Supino, 1897)
R. humeralis Rondelli, 1926
R. hurti Wilson, 1954
R. interventus Walker, Pegram and Keirans, 1995
R. jeanneli Neumann, 1913
R. kochi Dnitz, 1905
52
Table 1. (continued)

R. leporis Pomerantsev, 1946

R. longiceps Warburton, 1912
R. longicoxatus Neumann, 1905
R. longus Neumann, 1907
R. lounsburyi Walker, 1990
R. lunulatus Neumann, 1907
R. maculatus Neumann, 1901
R. masseyi Nuttall and Warburton, 1908
R. moucheti Morel, 1965
R. muehlensi Zumpt, 1943
R. muhsamae Morel and Vassiliades, 1965
R. neumanni Walker, 1990
R. nitens Neumann, 1904
R. oculatus Neumann, 1901
R. oreotragi Walker and Horak, 2000
R. pilans Schulze, 1935
R. planus Neumann, 1907
R. praetextatus Gerstcker, 1873
R. pravus Dnitz, 1910
R. pseudolongus Santos Dias, 1953
R. pulchellus (Gerstcker, 1873)
R. pumilio Schulze, 1935
R. punctatus Warburton, 1912
R. pusillus Gil Collado, 1936
R. ramachandrai Dhanda, 1966
R. rossicus Yakimov and Kol-Yakimova, 1911
R. sanguineus (Latreille, 1806)
R. scalpturatus Santos Dias, 1959
R. schulzei Olenev, 1929
R. sculptus Warburton, 1912
R. senegalensis Koch, 1844
R. serranoi Santos Dias, 1950
R. simpsoni Nuttall, 1910
R. simus Koch, 1844
R. sulcatus Neumann, 1908
R. supertritus Neumann, 1907
R. theileri Bedford and Hewitt, 1925
R. tricuspis Dnitz, 1906
R. turanicus Pomerantsev, 1936
 53
Table 1. (continued)

R. warburtoni Walker and Horak, 2000

R. zambeziensis Walker, Norval and Corwin, 1981
R. ziemanni Neumann, 1904
R. zumpti Santos Dias, 1950

Nuttalliellidae
Nuttalliellinae
Nuttalliella
N. namaqua Bedford, 1931

Acknowledgements

Meetings and discussions between the authors were made possible through
assistance received from the International Consortium on Ticks and Tick-
borne Diseases (ICTTD-2) project, financially supported by the INCO-DEV
program of the European Union under contract number ICA4-CT-2000-30006.
We wish to express our sincere thanks to Frans Jongejan, who coordinates the
ICTTD project. This research was also supported by the National Institute of
Allergy and Infectious Diseases grant AI 40729 to JEK.

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