Haemaphysalis

Haemaphysalis Temporal range: Cretaceous–present PreꞒ Ꞓ O S D C P T J K Pg N
Haemaphysalis longicornis
Scientific classification
Kingdom:	Animalia
Phylum:	Arthropoda
Subphylum:	Chelicerata
Class:	Arachnida
Order:	Ixodida
Family:	Ixodidae
Subfamily:	Haemaphysalinae
Genus:	Haemaphysalis Koch, 1844
Species
170 extant, see text. [1]

Haemaphysalis is a genus of hard ticks. ^[2] It is the second largest genus in the family Ixodidae, after Ixodes , and the largest genus of the Metastriata. Haemaphysalis ticks are found on all continents and across all major zoogeographical regions, except Antarctica, although most species are found in the Oriental and Afrotropical regions. Approximately 80% of the species are present in Asia and Africa, with significantly less species diversity in Europe and the Americas. ^[3] Haemaphysalis species primarily parasitise birds and mammals. ^[4]

Top down, L>R: Haemaphysalis longicornis , Haemaphysalis leporispalustris , Haemaphysalis pentalagi , Haemaphysalis bancrofti .

Top down, L>R: Haemaphysalis longicornis , Haemaphysalis leporispalustris , Haemaphysalis elliptica , Haemaphysalis sp., Haemaphysalis sp., Haemaphysalis bancrofti .

Description

Haemaphysalis are very small, elliptical ticks. They are best distinguished by the significant lateral salience on palpal article II, which extends beyond the margin of the basis capitulum. They are eyeless, with short mouthparts. ^[5]

Taxonomy

Haemaphysalis contains 170 species ^[6] distributed across 8 subgenera (Aboimisalis, Aborphysalis, Allophysalis, Haemaphysalis, Herpetobia, Kaiseriana, Ornithophysalis, Segalia) although the phylogeny of the genus is unresolved, with most subgenera being polyphyletic. ^{[7] [8]} The former subgenera Alloceraea and Sharifiella have been recently elevated to full generic genus rank. ^{[9] [10]}

1. Haemaphysalis aborensis Warburton, 1913
2. Haemaphysalis aciculifer Warburton, 1913
3. Haemaphysalis aculeata Lavarra, 1904
4. Haemaphysalis adleri Feldman-Muhsam, 1951
5. Haemaphysalis anomala Warburton, 1913
6. Haemaphysalis anomaloceraea Teng, 1984 ^[a]
7. Haemaphysalis anoplos Hoogstraal, Uilenberg & Klein, 1967
8. Haemaphysalis asiatica (Supino, 1897)
9. Haemaphysalis atheruri Hoogstraal, Trapido & Kohls, 1965
10. Haemaphysalis bancrofti Nuttall & Warburton, 1915
11. Haemaphysalis bandicota Hoogstraal & Kohls, 1965
12. Haemaphysalis bartelsi Schulze, 1938
13. Haemaphysalis bequaerti Hoogstraal, 1956
14. Haemaphysalis birmaniae Supino, 1897
15. Haemaphysalis bispinosa Neumann, 1897
16. Haemaphysalis bochkovi Apanaskevich & Tomlinson, 2019 ^[11]
17. Haemaphysalis borneata Hoogstraal, 1971
18. Haemaphysalis bremneri Roberts, 1963
19. Haemaphysalis burkinae Apanaskevich & Tomlinson, 2019 ^[11]
20. Haemaphysalis calcarata Neumann, 1902
21. Haemaphysalis calva Nuttall & Warburton, 1915
22. Haemaphysalis camicasi Tomlinson & Apanaskevich, 2019
23. Haemaphysalis campanulata Warburton, 1908
24. Haemaphysalis canestrinii (Supino, 1897)
25. Haemaphysalis capricornis Hoogstraal, 1966
26. Haemaphysalis caucasica Olenev, 1928
27. Haemaphysalis celebensis Hoogstraal, Trapido & Kohls, 1965
28. Haemaphysalis chordeilis (Packard, 1869)
29. Haemaphysalis cinnabarina Koch, 1844 ^[b]
30. Haemaphysalis colesbergensis Apanaskevich & Horak, 2008
31. Haemaphysalis concinna Koch, 1844
32. Haemaphysalis cooleyi Bedford, 1929
33. Haemaphysalis cornigera Neumann, 1897 ^[c]
34. Haemaphysalis cornupunctata Hoogstraal & Varma, 1962
35. Haemaphysalis cuspidata Warburton, 1910
36. Haemaphysalis dangi Phan Trong, 1977
37. Haemaphysalis danieli Černý & Hoogstraal, 1977
38. Haemaphysalis darjeeling Hoogstraal & Dhanda, 1970
39. Haemaphysalis davisi Hoogstraal, Dhanda & Bhat, 1970
40. Haemaphysalis demidovae Emel'yanova, 1978
41. Haemaphysalis doenitzi Warburton & Nuttall, 1909
42. Haemaphysalis elliptica (Koch, 1844)
43. Haemaphysalis elongata Neumann, 1897
44. Haemaphysalis erinacei Pavesi, 1884
45. Haemaphysalis eupleres Hoogstraal, Kohls & Trapido, 1965
46. Haemaphysalis filippovae Bolotin, 1979 ^[d]
47. Haemaphysalis flava Neumann, 1897
48. Haemaphysalis formosensis Neumann, 1913
49. Haemaphysalis fossae Hoogstraal, 1953
50. Haemaphysalis fujisana Kitaoka, 1970
51. Haemaphysalis galidiae Apanaskevich & Goodman, 2020 ^[12]
52. Haemaphysalis garhwalensis Dhanda & Bhat, 1968
53. Haemaphysalis mariae Apanaskevich, 2024 ^[13]
54. Haemaphysalis goral Hoogstraal, 1970
55. Haemaphysalis grochovskajae Kolonin, 1992
56. Haemaphysalis heinrichi Schulze, 1939
57. Haemaphysalis hirsuta Hoogstraal, Trapido & Kohls, 1966
58. Haemaphysalis hispanica Gil Collado, 1938
59. Haemaphysalis hoodi Warburton & Nuttall, 1909
60. Haemaphysalis hoogstraali Kohls, 1950
61. Haemaphysalis horaki Apanaskevich & Tomlinson, 2019 ^[11]
62. Haemaphysalis houyi Nuttall & Warburton, 1915
63. Haemaphysalis howletti Warburton, 1913
64. Haemaphysalis humerosa Warburton & Nuttall, 1909
65. Haemaphysalis hylobatis Schulze, 1933
66. Haemaphysalis hyracophila Hoogstraal, Walker & Neitz, 1971
67. Haemaphysalis hystricis Supino, 1897
68. Haemaphysalis indica Warburton, 1910
69. Haemaphysalis indoflava Dhanda & Bhat, 1968
70. Haemaphysalis intermedia Warburton & Nuttall, 1909
71. Haemaphysalis japonica Warburton, 1908
72. Haemaphysalis juxtakochi Cooley, 1946
73. Haemaphysalis kadarsani Hoogstraal & Wassef, 1977
74. Haemaphysalis kashmirensis Hoogstraal & Varma, 1962
75. Haemaphysalis kinneari Warburton, 1913
76. Haemaphysalis knobigera Prakasan & Ramani, 2007 ^[e]
77. Haemaphysalis koningsbergeri Warburton & Nuttall, 1909
78. Haemaphysalis kopetdaghica Kerbabaev, 1962
79. Haemaphysalis kumaonensis Geevarghese & Mishra, 2011
80. Haemaphysalis kutchensis Hoogstraal & Trapido, 1963
81. Haemaphysalis kyasanurensis Trapido, Hoogstraal & Rajagopalan, 1964
82. Haemaphysalis lagostrophi Roberts, 1963
83. Haemaphysalis lagrangei Larrousse, 1925
84. Haemaphysalis laocayensis Phan Trong, 1977
85. Haemaphysalis leachi (Audouin, 1826)
86. Haemaphysalis lemuris Hoogstraal, 1953
87. Haemaphysalis leporispalustris (Packard, 1869)
88. Haemaphysalis lobachovi Kolonin, 1995
89. Haemaphysalis longicornis Neumann, 1901
90. Haemaphysalis luzonensis Hoogstraal & Parrish, 1968
91. Haemaphysalis madagascariensis Colas-Belcour & Millot, 1948
92. Haemaphysalis mageshimaensis Saito & Hoogstraal, 1973
93. Haemaphysalis megalaimae Rajagopalan, 1963
94. Haemaphysalis megaspinosa Saito, 1969
95. Haemaphysalis menglaensis Pang, Chen & Xiang, 1982
96. Haemaphysalis minuta Kohls, 1950
97. Haemaphysalis mjoebergi Warburton, 1926
98. Haemaphysalis montgomeryi Nuttall, 1912
99. Haemaphysalis moreli Camicas, Hoogstraal & El Kammah, 1972
100. Haemaphysalis moschisuga Teng, 1980
101. Haemaphysalis muhsamae Santos Dias, 1954
102. Haemaphysalis nadchatrami Hoogstraal, Trapido & Kohls, 1965
103. Haemaphysalis nepalensis Hoogstraal, 1962
104. Haemaphysalis nesomys Hoogstraal, Uilenberg & Klein, 1966
105. Haemaphysalis norvali Hoogstraal & Wassef, 1983
106. Haemaphysalis novaeguineae Hirst, 1914
107. Haemaphysalis obesa Larrousse, 1925
108. Haemaphysalis obtusa Dönitz, 1910
109. Haemaphysalis oliveri Apanaskevich & Horak, 2008
110. Haemaphysalis orientalis Nuttall & Warburton, 1915
111. Haemaphysalis ornithophila Hoogstraal & Kohls, 1959
112. Haemaphysalis palawanensis Kohls, 1950
113. Haemaphysalis papuana Thorell, 1883
114. Haemaphysalis paraleachi Camicas, Hoogstraal & El Kammah, 1983
115. Haemaphysalis paraturturis Hoogstraal, Trapido & Rebello, 1963
116. Haemaphysalis parmata Neumann, 1905
117. Haemaphysalis parva Neumann, 1897
118. Haemaphysalis pavlovskyi Pospelova-Shtrom, 1935
119. Haemaphysalis pedetes Hoogstraal, 1972
120. Haemaphysalis pentalagi Pospelova-Shtrom, 1935
121. Haemaphysalis petrogalis Roberts, 1970
122. Haemaphysalis phasiana Saito, Hoogstraal & Wassef, 1974 ^[f]
123. Haemaphysalis pospelovashtromae Hoogstraal, 1966
124. Haemaphysalis princeps Tomlinson & Apanaskevich, 2019
125. Haemaphysalis psalistos Hoogstraal, Kohls & Parrish, 1967
126. Haemaphysalis punctaleachi Camicas, Hoogstraal & El Kammah, 1973
127. Haemaphysalis punctata Canestrini & Fanzago, 1878
128. Haemaphysalis qinghaiensis Teng, 1980
129. Haemaphysalis quadriaculeata Kolonin, 1992
130. Haemaphysalis ramachandrai Dhanda, Hoogstraal & Bhat, 1970
131. Haemaphysalis ratti Kohls, 1948
132. Haemaphysalis renschi Schulze, 1933
133. Haemaphysalis roubaudi Toumanoff, 1940
134. Haemaphysalis rugosa Santos Dias, 1956
135. Haemaphysalis rusae Kohls, 1950
136. Haemaphysalis sambar Hoogstraal, 1971
137. Haemaphysalis sciuri Kohls, 1950
138. Haemaphysalis semermis Neumann, 1901
139. Haemaphysalis shimoga Trapido & Hoogstraal, 1964 ^[g]
140. Haemaphysalis silacea Robinson, 1912
141. Haemaphysalis silvafelis Hoogstraal & Trapido, 1963
142. Haemaphysalis simplex Neumann, 1897
143. Haemaphysalis simplicima Hoogstraal & Wassef, 1979
144. Haemaphysalis sinensis Zhang, 1981
145. Haemaphysalis spinigera Neumann, 1897
146. Haemaphysalis spinulosa Neumann, 1906
147. Haemaphysalis subelongata Hoogstraal, 1953
148. Haemaphysalis subterra Hoogstraal, El Kammah & Camicas, 1992
149. Haemaphysalis sulcata Canestrini & Fanzago, 1878 ^[h]
150. Haemaphysalis sumatraensis Hoogstraal, El Kammah, Kadarsan & Anastos, 1971
151. Haemaphysalis sundrai Sharif, 1928
152. Haemaphysalis suntzovi Kolonin, 1993
153. Haemaphysalis susphilippensis Hoogstraal, Kohls & Parrish, 1968
154. Haemaphysalis taiwana Sugimoto, 1936
155. Haemaphysalis tauffliebi Morel, 1965
156. Haemaphysalis tibetensis Hoogstraal, 1965
157. Haemaphysalis tiptoni Hoogstraal, 1953
158. Haemaphysalis toxopei Warburton, 1927 ^[i]
159. Haemaphysalis traguli Oudemans, 1928
160. Haemaphysalis traubi Kohls, 1955
161. Haemaphysalis turturis Nuttall & Warburton, 1915
162. Haemaphysalis vasilisae Apanaskevich, 2025 ^[14]
163. Haemaphysalis verticalis Itagaki, Noda & Yamaguchi, 1944
164. Haemaphysalis vidua Warburton & Nuttall, 1909
165. Haemaphysalis vietnamensis Hoogstraal & Wilson, 1966
166. Haemaphysalis walkerae Apanaskevich & Tomlinson, 2019 ^[11]
167. Haemaphysalis warburtoni Nuttall, 1912
168. Haemaphysalis wellingtoni Nuttall & Warburton, 1908
169. Haemaphysalis yeni Toumanoff, 1944
170. Haemaphysalis zumpti Hoogstraal & El Kammah, 1974

Notes

1. Camicas et al. (1998) list as synonym of H. shimoga , while Kolonin (2009) treats both as synonyms of H. taiwana
2. This taxon is dubious - it has been extensively confused and synonymised with Haemaphysalis chordeilis and Haemaphysalis punctata . Its hosts are unknown. Most redescriptions are unclear, mistake the species for another, or misname the species as H. cinnaberina [sic].
3. This species is known in Japanese literature as Haemaphysalis ias Nakamura & Yajima, 1937
4. Possibly abnormal form of H. concinna (Kolonin 2009)
5. Taxon is ill-defined, and only considered provisionally valid.
6. Possible synonym of H. doenitzi (Kolonin, 2009)
7. Treated as synonym of H. taiwana in Kolonin (2009)
8. This taxon seems to have highly variable morphology, making it difficult to identify. Different authors depict profound differences in morphology across redescriptions, suggesting it may in fact be a species complex.
9. Kolonin (2009) treats as a synonym of H. kinneari

References

1. Guglielmone, Alberto A; Petney, Trevor N; Robbins, Richard G (2020-11-05). "Ixodidae (Acari: Ixodoidea): descriptions and redescriptions of all known species from 1758 to December 31, 2019" . Zootaxa (Monograph). 4871 (1). Auckland: Magnolia Press: 1–322. doi:10.11646/zootaxa.4871.1.1. eISSN   1175-5326. ISBN   978-1-77688-088-1. OCLC   8837433568. PMID   33311340.
2. Koch, C.L. (1844) Systematische Übersicht über die Ordnung der Zecken. Archiv für Naturgeschichte, 10, 217–239. https://doi.org/10.5962/bhl.part.29560
3. Guglielmone, Alberto A.; Nava, Santiago; Robbins, Richard G. (2021). "Neotropical Hard Ticks (Acari: Ixodida: Ixodidae)". SpringerLink. doi:10.1007/978-3-030-72353-8.
4. Mehlhorn, Heinz (2016), "Haemaphysalis Species", Encyclopedia of Parasitology, Springer, Berlin, Heidelberg, pp. 1198–1200, doi:10.1007/978-3-662-43978-4_1366, ISBN   978-3-662-43978-4 , retrieved 2025-10-05
5. Barker, Stephen; Barker, Dayana (2023). Ticks of Australasia: 125 species of ticks in and around Australia. Auckland, New Zealand: Zootaxa, Magnolia Press. p. 12. ISBN   978-1-77688-700-2.
6. Guglielmone, Alberto A; Petney, Trevor N; Robbins, Richard G (2020-11-05). "Ixodidae (Acari: Ixodoidea): descriptions and redescriptions of all known species from 1758 to December 31, 2019" . Zootaxa (Monograph). 4871 (1). Auckland: Magnolia Press: 1–322. doi:10.11646/zootaxa.4871.1.1. eISSN   1175-5326. ISBN   978-1-77688-088-1. OCLC   8837433568. PMID   33311340.
7. Kelava, Samuel; Apanaskevich, Dmitry A.; Shao, Renfu; Gofton, Alexander W.; Mans, Ben J.; Teo, Ernest J. M.; Norval, Gerrut; Barker, Dayana; Nakao, Ryo; Barker, Stephen C. (June 2024). "Insights from entire mitochondrial genome sequences into the phylogeny of ticks of the genera Haemaphysalis and Archaeocroton with the elevation of the subgenus Alloceraea Schulze, 1919 back to the status of a genus". Medical and Veterinary Entomology. 38 (2): 189–204. doi:10.1111/mve.12708. ISSN   0269-283X.
8. Kelava, Samuel; Mans, Ben J.; Shao, Renfu; Barker, Dayana; Teo, Ernest J.M.; Chatanga, Elisha; Gofton, Alexander W.; Moustafa, Mohamed Abdallah Mohamed; Nakao, Ryo; Barker, Stephen C. (March 2023). "Seventy-eight entire mitochondrial genomes and nuclear rRNA genes provide insight into the phylogeny of the hard ticks, particularly the Haemaphysalis species, Africaniella transversale and Robertsicus elaphensis". Ticks and Tick-borne Diseases. 14 (2) 102070. doi:10.1016/j.ttbdis.2022.102070.
9. Kelava, Samuel; Apanaskevich, Dmitry A.; Shao, Renfu; Gofton, Alexander W.; Mans, Ben J.; Teo, Ernest J. M.; Norval, Gerrut; Barker, Dayana; Nakao, Ryo; Barker, Stephen C. (2024). "Insights from entire mitochondrial genome sequences into the phylogeny of ticks of the genera Haemaphysalis and Archaeocroton with the elevation of the subgenus Alloceraea Schulze, 1919 back to the status of a genus". Medical and Veterinary Entomology. 38 (2): 189–204. doi:10.1111/mve.12708. ISSN   0269-283X.
10. Apanaskevich, Dmitry A.; Greiman, Stephen E.; Goodman, Steven M.; Apanaskevich, Maria A.; Ahmed, Rokeya; Barker, Stephen C. (2025-04-29). "Validation of Sharifiella Santos Dias, 1958 (Acari: Ixodidae) as a genus of hard ticks containing S. theilerae (Hoogstraal, 1953), a parasite of tail-less tenrec, Tenrec ecaudatus (Schreber) (Afrosoricida: Tenrecidae) on Madagascar". Zootaxa. 5627 (3): 503–525. doi:10.11646/zootaxa.5627.3.5. ISSN   1175-5334.
11. Dmitry Apanaskevich and Jackson A. Tomlinson. 2019. Description of four new species of Haemaphysalis Koch, 1844 (Acari: Ixodidae) from the H. (Rhipistoma) spinulosa subgroup, parasites of carnivores and rodents in Africa. Systematic Parasitology , July, First Online: 22 August 2019. DOI: 10.1007/s11230-019-09875-7, https://link.springer.com/article/10.1007/s11230-019-09875-7.
12. Dmitry Apanaskevich and Steven M. Goodman. 2020. Description of a new species of Haemaphysalis Koch, 1844 (Acari: Ixodidae) from the H. (Rhipistoma) asiatica subgroup, parasite of an endemic Malagasy carnivoran (Carnivora: Eupleridae). Systematic Parasitology, 97: 591–599; https://link.springer.com/article/10.1007/s11230-020-09943-3, DOI: 10.1007/s11230-020-09943-3.
13. Apanaskevich, Dmitry A. (2024-07-26). "Description of a new species of Haemaphysalis Koch, 1844 (Acari: Ixodidae), a parasite of hares and rabbits (Lagomorpha: Leporidae) in Texas, Oklahoma and Colorado (USA), that was misidentified as H. leporispalustris (Packard, 1869) for more than a century". Zootaxa. 5486 (3): 435–450. doi:10.11646/zootaxa.5486.3.6. ISSN   1175-5334.
14. Apanaskevich, Dmitry A. (2025-09-08). "Description of a new species of Haemaphysalis Koch, 1844 (Acari: Ixodidae), an ornate parasite of birds in Florida, South Carolina and Texas (USA), and redescription of all parasitic stages of H. chordeilis (Packard, 1869)". Zootaxa. 5691 (1): 45–74. doi:10.11646/zootaxa.5691.1.2. ISSN   1175-5334.