Comments and Conclusions

Home , Amblyomma cajennense, Amblyomma testudinarium, Amblyomma triguttatum, Cosmiomma, Dermacentor, Dermacentor albipictus

At least 283 species of Ixodidae have been recorded as parasites of humans, representing 38.8% of all taxa in this family. A total of 69 species of Ixodes (Prostriata) have been recovered from people, while 214 species in all genera of the Metastriata have been found on humans, representing 27.3% and 45.0% of taxa in these groups, respectively. Numerically, the most important genera causing human parasitism are Ixodes (69 species), Amblyomma (63) and Haemaphysalis (60), but the genera with the greatest percentages of species attacking humans are Hyalomma (66.7%), Rhipicephalus (56.5%) and Dermacentor (55.0%) (Table 1). Table 2 lists all tick species that have been found on people, their distribution by zoogeographic region(s), and, for those species that occur in more than one region,

Table 1 Numbers and percentages of ixodid species found feeding on humans, by genus. N° species feeding on humans Genus Total N° species (%) (% of each genus) Ixodes 253 (34.7) 69 (27.3) Haemaphysalis 166 (22.8) 60 (36.1) Amblyomma 138 (18.9) 63 (45.7) Rhipicephalus 85 (11.7) 48 (56.5) Dermacentor 40 (5.5) 22 (55.0) Hyalomma 27 (3.7) 18 (66.7) Bothriocroton 7 (1.0) 2 (28.6) Anomalohimalaya 3 (0.4) 0 (0.0) Margaropus 3 (0.4) 0 (0.0) Nosomma 2 (0.3) 1 (50.0) Rhipicentor 2 (0.3) 0 (0.0) Compluriscutula (fossil) 1 (0.1) 0 (0.0) Cornupalpatum (fossil) 1 (0.1) 0 (0.0) Cosmiomma 1 (0.1) 0 (0.0) Total 729 283 (38.8)

© Springer International Publishing AG, part of Springer Nature 2018 231 A. A. Guglielmone, R. G. Robbins, Hard Ticks (Acari: Ixodida: Ixodidae) Parasitizing Humans, https://doi.org/10.1007/978-3-319-95552-0 232 Comments and Conclusions

Table 2 Species of Ixodidae found on humans, by zoogeographic region, frequency of parasitism, and tick stages involved. The most common region and most common tick stage(s) found on humans are underlined. M: males, F: females, N: nymphs, L: larvae, A: adults but sex unknown, I: immature stages without further definition, U: unknown. Tick species Zoogeographic region (s) Frequency Tick stage(s) Ixodes acuminatus Palearctic Sporadic M, F, N I. acutitarsus Oriental, Palearctic Frequent M, F, L I. angustus Nearctic, Palearctic Sporadic F, N I. apronophorus Palearctic Very rare A I. asanumai Oriental, Palearctic Very rare F I. australiensis Australasian Very rare M I. baergi Nearctic Very rare U I. banksi Nearctic Rare M I. boliviensis Neotropical Rare M, F I. brunneus Nearctic, Neotropical Very rare F I. canisuga Palearctic Very rare U I. cavipalpus Afrotropical Very rare M I. columnae Palearctic Very rare L I. confusus Australasian Very rare F I. cookei Nearctic, Neotropical Very frequent F, N, L I. cornuatus Australasian Sporadic U I. crenulatus Palearctic Very rare U I. cumulatimpunctatus Afrotropical Rare F I. dentatus Nearctic Sporadic N, L I. eichhorni Australasian, Oriental Very rare F I. fecialis Australasian Very rare U I. festai Palearctic Very rare A I. frontalis Palearctic Very rare F I. gibbosus Palearctic Sporadic F, N I. granulatus Australasian, Oriental, Palearctic Rare F I. hexagonus Palearctic Frequent M, F, N., L I. holocyclus Australasian Frequent M, F, N, L I. kashmiricus Oriental, Palearctic Rare F, N I. kazakstani Palearctic Very rare U I. kingi Nearctic Rare F, N I. kohlsi Australasian Very rare N I. laguri Palearctic Sporadic F, N, L I. marxi Nearctic Frequent F, N I. monospinosus Palearctic Sporadic M, F, N I. muniensis Afrotropical Very rare N I. muris Nearctic Frequent F, N, L I. myrmecobii Australasian Very rare F I. nipponensis Oriental, Palearctic Frequent F, N I. ovatus Oriental, Palearctic Frequent M, F, N, L (continued) Comments and Conclusions 233

Table 2 (continued) Tick species Zoogeographic region (s) Frequency Tick stage(s) I. pacificus Nearctic Frequent M, F, N, L I. pararicinus Neotropical Very rare N I. pavlovskyi Palearctic Frequent F I. persulcatus Oriental, Palearctic Very frequent M, F, N, L I. petauristae Oriental Very rare N, L I. pilosus Afrotropical Sporadic F, N I. rageaui Afrotropical Very rare M I. rasus Afrotropical Very rare F I. redikorzevi Palearctic Sporadic F I. ricinus Palearctic Very frequent M, F, N, L I. rubicundus Afrotropical Very rare F I. rugosus Nearctic Very rare A I. scapularis Nearctic, Neotropical Very frequent M, F, N, L I. schillingsi Afrotropical Very rare F I. sculptus Nearctic Rare A, N I. sinensis Oriental Very rare U I. soricis Nearctic Very rare F I. spinicoxalis Oriental Very rare N I. spinipalpis Nearctic, Neotropical Sporadic F, N I. tancitarius Nearctic Very rare F I. tanuki Oriental, Palearctic Very rare F I. tasmani Australasian Sporadic F, N I. texanus Nearctic Rare N, L I. trianguliceps Palearctic Very rare U I. turdus Palearctic Rare F I. uriae Circumpolar Frequent F, N I. vanidicus Afrotropical Very rare M, F I. ventalloi Palearctic Rare F I. vespertilionis Afrotropical, Oriental, Palearctic Very rare N I. woodi Nearctic Very rare N Amblyomma americanum Nearctic Very frequent M, F, N, L A. aureolatum Neotropical Sporadic M, F A. babirussae Australasian Sporadic M, F, L A. brasiliense Neotropical Sporadic M, F, N, L A. breviscutatum Australasian, Oriental Sporadic M, F A. cajennense Neotropical Frequent M, F, N, L A. calcaratum Neotropical Very rare U A. coelebs Nearctic, Neotropical Frequent M, F, N, L A. cohaerens Afrotropical Very rare U A. cordiferum Australasian, Oriental Very rare N A. dissimile Nearctic, Neotropical Sporadic F, N A. dubitatum Neotropical Sporadic M, F, N, L (continued) 234 Comments and Conclusions

Table 2 (continued) Tick species Zoogeographic region (s) Frequency Tick stage(s) A. falsomarmoreum Afrotropical Very rare F A. fuscum Neotropical Very rare F A. gemma Afrotropical Very rare M A. geoemydae Oriental, Palearctic Very rare F, N A. hadanii Neotropical Sporadic F, N, L A. hebraeum Afrotropical Frequent M, F, N, L A. incisum Neotropical Sporadic F, N A. inornatum Nearctic, Neotropical Rare M A. integrum Oriental Frequent F, N A. javanense Oriental Very rare U A. latepunctatum Neotropical Very rare M, F A. latum Afrotropical Very rare A A. lepidum Afrotropical, Palearctic Rare M, F A. limbatum Australasian Rare U A. loculosum Afrotropical, Australasian, Oriental* Sporadic M, F, N A. longirostre Neotropical Rare M, F, N A. maculatum Nearctic, Neotropical Frequent M, F, N, L A. marmoreum Afrotropical Sporadic M, F, N, L A. mixtum Nearctic, Neotropical Frequent M, F, N, L A. moreliae Australasian Very rare U A. naponense Neotropical Sporadic M, F, N, L A. neumanni Neotropical Very frequent M, F, N, L A. nuttalli Afrotropical Sporadic M, N, L A. oblongoguttatum Nearctic, Neotropical Sporadic M, F, N A. ovale Nearctic, Neotropical Frequent M, F A. pacae Neotropical Very rare M A. parkeri Neotropical Very rare N A. parvum Neotropical Frequent M, F, N A. paulopunctatum Afrotropical Very rare M, N A. pecarium Neotropical Rare N A. personatum Afrotropical Very rare A A. postoculatum Australasian Very rare F A. pseudoconcolor Neotropical Very rare U A. pseudoparvum Neotropical Rare M, F A. romitii Neotropical Rare M, L A. rotundatum Nearctic, Neotropical, Pacific Island Very rare F A. sabanerae Neotropical Very rare A A. scalpturatum Neotropical Sporadic M, F, N A. sculptum Neotropical Frequent M, F, N, L A. sparsum Afrotropical Very rare U A. tapirellum Neotropical Sporadic A, N A. tenellum Nearctic, Neotropical Sporadic M, F, N, L (continued) Comments and Conclusions 235

Table 2 (continued) Tick species Zoogeographic region (s) Frequency Tick stage(s) A. testudinarium Australasian, Oriental, Palearctic Very frequent M, F, N, L A. tholloni Afrotropical Sporadic M, F, N, L A. tigrinum Neotropical Sporadic M, F, N A. tonelliae Neotropical Sporadic M, F, N, L A. triguttatum Australasian Frequent M, F, N, L A. triste Nearctic, Neotropical Sporadic M, F A. tuberculatum Nearctic Frequent L A. variegatum Afrotropical, Neotropical Frequent M, F, L A. varium Neotropical Very rare N Bothriocroton auruginans Australasian Very rare A B. hydrosauri Australasian Rare U Dermacentor albipictus Nearctic, Neotropical Frequent M, F, N, L D. andersoni Nearctic Very frequent M, F, N, L D. atrosignatus Australasian, Oriental Sporadic M, F D. auratus Oriental Frequent M, F, N, L D. bellulus Oriental, Palearctic Very rare M D. circumguttatus Afrotropical Rare M, F D. compactus Oriental Rare M, F D. hunteri Nearctic Sporadic M, F D. imitans Neotropical Very rare M, F D. latus Neotropical Very rare M, F D. marginatus Palearctic Very frequent M, F, N D. nitens Nearctic, Neotropical Sporadic M, F, N, L D. niveus Palearctic Sporadic M, F D. nuttalli Palearctic Very frequent M, F, N D. occidentalis Nearctic Frequent M, F, N, L D. parumapertus Nearctic Rare M, F, N D. raskemensis Oriental, Palearctic Rare M D. reticulatus Palearctic Frequent M, F, N D. rhinocerinus Afrotropical Very rare A D. silvarum Palearctic Very frequent M, F, N D. steini Australasian, Oriental Sporadic M, F D. variabilis Nearctic, Neotropical Very frequent M, F, N, L Haemaphysalis aculeata Oriental Rare M, F, N H. anomala Oriental Rare M, F H. aponommoides Oriental, Palearctic Sporadic M, F, N H. bancrofti Australasian, Oriental Sporadic F, N H. birmaniae Oriental Very rare N H. bispinosa Oriental Rare F, N H. campanulata Oriental, Palearctic Rare F H. caucasica Palearctic Very rare F H. celebensis Australasian Very rare F (continued) 236 Comments and Conclusions

Table 2 (continued) Tick species Zoogeographic region (s) Frequency Tick stage(s) H. chordeilis Nearctic Rare M, F H. colasbelcouri Oriental Very rare M H. concinna Oriental, Palearctic Frequent M, F, N H. cornigera Australasian, Oriental, Palearctic Very rare F H. cuspidata Oriental Very rare U H. darjeeling Oriental Very rare M H. doenitzi Australasian, Oriental, Palearctic Very rare N H. elliptica Afrotropical Sporadic M, F, L H. elongata Afrotropical Very rare N H. erinacei Palearctic Rare M, F H. flava Oriental, Palearctic Sporadic M, F, N H. heinrichi Oriental Very rare M H. hirsuta Oriental Sporadic A, I H. hoodi Afrotropical Very rare A H. humerosa Australasian Very rare U H. hylobatis Oriental Very rare A H. hystricis Australasian, Oriental, Palearctic Sporadic M, F, N H. indoflava Oriental Very rare U H. inermis Palearctic Sporadic A H. intermedia Oriental Very rare U H. japonica Palearctic Sporadic M, F, N, L H. juxtakochi Nearctic, Neotropical Sporadic M, F, N H. kitaokai Oriental, Palearctic Very rare F H. koningsbergeri Oriental Very rare M H. lagrangei Oriental Rare M, F, N H. leachi Afrotropical, Palearctic Very rare M, F H. leporispalustris Nearctic, Neotropical Sporadic F, N H. longicornis Australasian, Oriental, Palearctic Frequent M, F, N, L H. mageshimaensis Oriental, Palearctic Sporadic M, F, N H. megaspinosa Palearctic Very rare F, N H. mjoebergi Oriental Very rare M, F H. montgomeryi Oriental, Palearctic Rare M, F H. nadchatrami Oriental Sporadic M, F H. nepalensis Oriental, Palearctic Very rare F, N H. novaeguineae Australasian Very rare M, N H. obesa Oriental Very rare A, N H. papuana Australasian, Oriental Very rare A H. paraleachi Afrotropical Very rare M H. parmata Afrotropical Very rare F, N H. parva Palearctic Very frequent M, F, N H. punctata Palearctic Frequent M, F, N H. qinghaiensis Oriental, Palearctic Very rare U (continued) Comments and Conclusions 237

Table 2 (continued) Tick species Zoogeographic region (s) Frequency Tick stage(s) H. ramachandrai Oriental Very rare F H. roubaudi Oriental Very rare M H. semermis Oriental Rare M, F, N H. shimoga Oriental Very rare F H. silacea Afrotropical Very rare N H. spinigera Oriental Very frequent N, L H. sulcata Afrotropical, Oriental, Palearctic Frequent M, F, N H. turturis Oriental Sporadic N, L H. wellingtoni Australasian, Oriental Very rare F Hyalomma aegyptium Palearctic Very frequent M, F, N H. albiparmatum Afrotropical Very rare M H. anatolicum Afrotropical, Oriental, Palearctic Frequent M, F H. asiaticum Palearctic Sporadic A, N H. brevipunctatum Oriental Sporadic M, F, N H. dromedarii Afrotropical, Oriental, Palearctic Rare M H. excavatum Afrotropical, Palearctic Frequent M, F H. glabrum Afrotropical Very rare A H. hussaini Oriental Very rare F H. impeltatum Afrotropical, Palearctic Rare M, N H. isaaci Oriental, Palearctic Sporadic M, F, N H. lusitanicum Palearctic Sporadic M, F H. marginatum Palearctic Very frequent M, F, N H. rufipes Afrotropical, Palearctic Sporadic M, F, L H. schulzei Palearctic Very rare F H. scupense Oriental, Palearctic Frequent M, F H. truncatum Afrotropical Sporadic M, F H. turanicum Afrotropical, Oriental, Palearctic Frequent M, F Nosomma monstrosum Oriental Very rare M Rhipicephalus annulatus Afrotropical, Nearctic, Palearctic Sporadic M, F R. appendiculatus Afrotropical Sporadic M, F, N, L R. armatus Afrotropical Very rare M, F R. aurantiacus Afrotropical Very rare M R. australis Australasian, Oriental, Pacific islands Very rare F, L R. bequaerti Afrotropical Sporadic M, F R. bursa Palearctic Very frequent M, F, N R. carnivoralis Afrotropical Very rare A R. complanatus Afrotropical Sporadic M, F R. compositus Afrotropical Very rare M R. decoloratus Afrotropical Rare M, F, L R. distinctus Afrotropical Very rare A, N R. evertsi Afrotropical Sporadic M, F, N, L R. follis Afrotropical Rare M, F (continued) 238 Comments and Conclusions

Table 2 (continued) Tick species Zoogeographic region (s) Frequency Tick stage(s) R. fulvus Afrotropical, Palearctic Very rare M R. gertrudae Afrotropical Sporadic M, F, N R. glabroscutatus Afrotropical Very rare M R. guilhoni Afrotropical Very rare M R. haemaphysaloides Australasian, Oriental, Palearctic Sporadic M, F, N R. humeralis Afrotropical Very rare A R. hurti Afrotropical Very rare A R. jeanneli Afrotropical Very rare A R. kochi Afrotropical Very rare F R. longus Afrotropical Very rare A R. lunulatus Afrotropical Rare MF R. maculatus Afrotropical Sporadic M, F, N R. microplus All regions Sporadic M, F, N, L R. muehlensis Afrotropical Very rare M R. muhsamae Afrotropical, Palearctic Rare M, F R. pilans Australasian, Oriental Very rare M R. planus Afrotropical Rare F R. praetextatus Afrotropical, Palearctic Rare F R. pravus Afrotropical Sporadic M, F R. pulchellus Afrotropical Sporadic M, F, N, L R. pumilio Oriental, Palearctic Sporadic M, F R. pusillus Palearctic Rare M, F R. rossicus Palearctic Sporadic A R. sanguineus sensu lato All regions Very frequent M, F, N, L R. schulzei Palearctic Rare A R. senegalensis Afrotropical Rare M, F R. simus Afrotropical Sporadic M, F R. sulcatus Afrotropical Rare M R. supertritus Afrotropical Very rare M R. turanicus sensu lato Afrotropical, Oriental, Palearctic Very frequent M, F, N R. warburtoni Afrotropical Very rare M, F R. zambeziensis Afrotropical Very rare N R. ziemanni Afrotropical Rare M, F R. zumpti Afrotropical Very rare A *Most records from islands near the Afrotropical, Australasian and Oriental mainland. the region where they are most often found. Table 2 also classifies these species according to the frequency with which they parasitize humans, and the tick stages reported from people. When known, the most common stages found on people are provided; however, in most cases, such information is unavailable, and in this table the dominant stages infesting humans are known for only 69 taxa (24% of the total found on humans). Stages found on people are unknown for 22 (8%) of the ticks causing human parasitism (nine Ixodes, seven Amblyomma, five Haemaphysalis, Comments and Conclusions 239 and one Bothriocroton); conversely, tick stages found on people are known for all species of Dermacentor, Hyalomma and Rhipicephalus. Although data are limited, it appears that in the genus Ixodes females are the predominant stage infesting humans, nymphs predominate in the genus Amblyomma, and both male and female ticks in Dermacentor, but stage predominance is unknown for ticks of other genera. The only genus in which males are the most common stage causing human parasitism is Hyalomma (three species), while larvae are the principal human parasites in five species (one Ixodes, one Dermacentor, and three Amblyomma). It seems likely that the priority of larvae, and perhaps nymphs, as parasites of humans has been underestimated because of the difficulties that often attend identification of these stages. Globally, the species of ticks that bite humans are found in the 20 combinations of zoogeographic regions shown in Table 3. However, 242 species (86% of the total) occur in just the first eight zoogeographic distributions listed in Table 3. As expected, the numbers and percentages of ticks causing human parasitism in these regions do not reflect the figures for all species found in these regions as compiled by Guglielmone et al. (2014), and new taxa described subsequently. Thus, in four zoogeographic regions the species of ticks known to feed on people are proportionally less than the species actually present: the Afrotropical Region contains 64 species that feed on humans, representing 23% of the total, but this region accounts for 29% of all ixodid taxa; the Oriental Region includes 33 species (12%) that feed on people, but 14% of all species; the Neotropical Region comprises 30 species (11%) known from people, but 14% of all ixodids; and the Australasian Region encom- passes 18 species (6%) that cause human parasitism, but 8% of all hard tick taxa. Conversely, four zoogeographic regions or combinations of regions contain numbers of tick species found on people that are equal to or higher than their proportions of the global hard tick fauna: the Palearctic Region contains 39 species that feed on humans, representing 14% of the total, and a matching 14% of all hard tick species; the Nearctic Region includes 20 species (7%) found on people, but only 5% of all species; the zoogeographic combination Oriental-Palearctic accounts for 22 species (8%), but only 5% of hard ticks worldwide share this distribution; and the combination Nearctic-Neotropical comprises 18 species (6%), but only 4% of hard tick taxa share this zoogeographical distribution. These apparently higher proportions involve a majority of the species found in the Nearctic and Palearctic Regions, indicating that either most ticks present in these regions are more prone to attack humans than species elsewhere, or that more tick studies have been conducted in these regions. A total of 117 (41%) hard tick species have very rarely been found on people, 69 (24%) sporadically, 45 (16%) rarely, 33 (12%) frequently, and 19 (7%) species have been collected from humans very frequently (Table 2). Table 4 shows the distribution of these frequencies according to the genera involved. The genera Amblyomma, Haemaphysalis, Ixodes and Rhipicephalus contain the highest numbers of species ranked as rare or very rare human parasites, with the percentage for these two categories reaching 68% in the case of genus Haemaphysalis. On the other hand, the genera Dermacentor and Hyalomma contain essentially the same numbers in the combinations very rare and rare as in the combinations very frequently and 240 Comments and Conclusions

Table 3 Numbers and percentages of ixodid species found feeding on humans by zoogeographic region(s) and frequency of parasitism. Zoogeographic region(s) N° (%) Frequency N° (%) Afrotropical 64 (23) Very rare 39 (61) Sporadic 15 (23) Rare 9 (14) Frequent 1 (2) Palearctic 40 (14) Very rare 12 (30) Sporadic 11 (28) Very frequent 8 (20) Rare 5 (13) Frequent 4 (10) Oriental 33 (12) Very rare 20 (61) Rare 6 (18) Sporadic 4 (12) Frequent 2 (6) Very frequent 1 (3) Neotropical 30 (11) Very rare 11 (37) Sporadic 10 (33) Rare 5 (17) Frequent 3 (10) Very frequent 1 (3) Oriental, Palearctic 22 (8) Very rare 7 (32) Frequent 5 (23) Sporadic 5 (23) Rare 4 (17) Very frequent 1 (5) Nearctic 20 (7) Rare 6 (30) Very rare 5 (25) Frequent 4 (20) Sporadic 3 (15) Very frequent 2 (10) Australasian 18 (6) Very rare 11 (61) Sporadic 3 (17) Frequent 2 (11) Rare 2 (11) Nearctic, Neotropical 18 (6) Sporadic 8 (44) Frequent 5 (28) Very frequent 3 (17) Rare 1 (8) Very rare 1 (6) Australasian, Oriental 9 (3) Very rare 5 (56) Sporadic 4 (44) (continued) Comments and Conclusions 241

Table 3 (continued) Zoogeographic region(s) N° (%) Frequency N° (%) Afrotropical, Palearctic 7 (2) Rare 4 (57) Frequent 1 (14) Sporadic 1 (14) Very rare 1 (14) Australasian, Oriental, Palearctic 7 (2) Sporadic 2 (29) Very rare 2 (29) Very frequent 1 (14) Frequent 1 (14) Rare 1 (14) Afrotropical, Oriental, Palearctic 6 (2) Frequent 3 (50) Very frequent 1 (17) Rare 1 (17) Very rare 1 (17) All Zoogeographic Regions 2 (1) Very frequent 1 (50) Sporadic 1 (50) Afrotropical, Australasian, Oriental* 1 (< 1) Sporadic Afrotropical, Nearctic, Palearctic 1 (< 1) Sporadic Afrotropical, Neotropical 1 (< 1) Frequent Australasian, Oriental, Pacific Islands 1 (< 1) Very rare Circumpolar 1 (< 1) Frequent Nearctic, Neotropical, Pacific Island 1 (< 1) Very rare Nearctic, Palearctic 1 (< 1) Sporadic *Most records from islands near the Afrotropical, Australasian and Oriental mainland. frequently on humans. The genera Dermacentor and Hyalomma are strongly represented in the Palearctic Zoogeographic Region, with 15 of the 40 species of Dermacentor found there, of which 12 are present only in this region, while 16 of the 27 species of Hyalomma are found in the Palearctic Region, seven of them exclusively so (Guglielmone et al. 2014, and new taxa described subsequently). This result is probably a consequence of investment in research on human tick- borne diseases in Europe and Palearctic Asia, although it cannot be ruled out that species in these genera are more prone to bite people than those in other genera. All ixodid ticks found on humans should be regarded as posing a risk to human health. Even those species known from only one or a few records from people (e.g., Amblyomma hadanii) may be of greater importance than such records suggest. Additionally, tick species that are rarely found on humans should alert epidemiolo- gists to the wide range of potential vectors that exist in nature and that we may expect to encounter in the future as human activities penetrate tick habitats that were previously undisturbed. Some ticks found sporadically on people may represent species whose capacity as parasites or disease vectors is underestimated due to limited research. Clearly, however, the 52 species that frequently or very frequently feed on humans are of concern, since many of them are already known to transport pathogens, and the geographic distributions of these species are shown in Table 5. 242 Comments and Conclusions

Table 4 Numbers of ixodid species found on humans, by genus, together with percentages of the total number of human-feeding species, and degrees of frequency of human parasitism. Genera N° (%) Frequency N° (%) Amblyomma 63 (22) Very frequent 3 (5) Frequent 12 (19) Sporadic 20 (32) Rare 7 (11) Very rare 20 (32) Bothriocroton 2 (<1) Rare 1 (50) Very rare 1 (50) Dermacentor 22 (8) Very frequent 5 (23) Frequent 4 (18) Sporadic 5 (23) Rare 4 (18) Very rare 4 (18 Haemaphysalis 60 (21) Very frequent 2 (3) Frequent 4 (7) Sporadic 13 (22) Rare 9 (15) Very rare 32 (53) Hyalomma 18 (6) Very frequent 2 (11) Frequent 4 (22) Sporadic 6 (33) Rare 2 (11) Very rare 4 (22) Ixodes 69 (24) Very frequent 4 (6) Frequent 9 (13) Sporadic 12 (17) Rare 10 (14) Very rare 33 (48) Nosomma 1 (<1) Very rare 1 (100) Rhipicephalus 48 (17) Very frequent* 3 (6) Sporadic 14 (29) Rare 11 (23) Very rare 20 (42) *Two of the three Rhipicephalus species listed as frequently found on humans in fact represent species complexes.

A total of 27 species found very frequently or frequently on people occur in just one zoogeographic region, while the remaining 25 frequent or very frequent species inhabit more than one region, with Rhipicephalus sanguineus s. l. recorded from all regions of the world (Table 5). Twenty-six of these species cause human parasitism in the Palearctic Region, and although Ixodes nipponensis, Haemaphysalis sulcata, Hyalomma excavatum and Hyalomma scupense are established in this and other regions, all documented cases of these ticks feeding on humans have been reported Comments and Conclusions 243

Table 5 Species of Ixodidae found very frequently or frequently on humans and their range, by zoogeographic region(s). The most common region is underlined. Very frequent Frequent Tick species Zoogeographic range Tick species Zoogeographic range Ixodes cookei Nearctic, Neotropical Ixodes acutitarsus Oriental, Palearctic I. persulcatus Oriental, Palearctic I. hexagonus Palearctic I. ricinus Palearctic I. holocyclus Australasian I. scapularis Nearctic, Neotropical I. muris Nearctic Amblyomma Nearctic I. nipponensis Oriental, Palearctic americanum A. neumanni Neotropical I. ovatus Oriental, Palearctic A. testudinarium Australasian, Oriental, I. pacificus Nearctic Palearctic Dermacentor Nearctic I. pavlovskyi Palearctic andersoni D. marginatus Palearctic I. uriae Circumpolar D. nuttalli Palearctic Amblyomma Neotropical cajennense D. silvarum Palearctic A. coelebs Nearctic, Neotropical D. variabilis Nearctic, Neotropical A. hebraeum Afrotropical Haemaphysalis Palearctic A. integrum Oriental parva H. spinigera Oriental A. maculatum Nearctic, Neotropical Hyalomma Palearctic A. mixtum Nearctic, Neotropical aegyptium H. marginatum Palearctic A. ovale Nearctic, Neotropical Rhipicephalus bursa Palearctic A. parvum Neotropical R. sanguineus sensu All regions A. sculptum Neotropical lato R. turanicus sensu Afrotropical, Oriental, A. triguttatum Australasian lato Palearctic A. tuberculatum Nearctic A. variegatum Afrotropical, Neotropical Dermacentor Nearctic, Neotropical albipictus D. auratus Oriental D. occidentalis Nearctic D. reticulatus Palearctic Haemaphysalis Oriental, Palearctic concinna H. longicornis Australasian, Oriental, Palearctic H. punctata Palearctic H. sulcata Afrotropical, Oriental, Palearctic (continued) 244 Comments and Conclusions

Table 5 (continued) Very frequent Frequent Tick species Zoogeographic range Tick species Zoogeographic range Hyalomma Afrotropical, Oriental, anatolicum Palearctic H. excavatum Afrotropical, Palearctic H. scupense Oriental, Palearctic H. turanicum Afrotropical, Oriental, Palearctic only from Palearctic localities. By contrast, a total of 16 species found frequently on people occur in the Oriental Region (exclusively or in combination with other regions), but no cases of human parasitism have been reported for Ixodes nipponensis, Haemaphysalis sulcata, Hyalomma excavatum and Hyalomma scupense in the Oriental Region. A situation similar to that in the Palearctic Region can be seen in the Nearctic, where 14 species that very frequently or frequently attack humans are present, several also occurring in the Neotropical Region, such as Ixodes cookei, Ixodes scapularis, Dermacentor albipictus and Dermacentor variabilis, but all records of human parasitism are exclusively from the Nearctic. This pattern is mir- rored in the Neotropics, for while Amblyomma coelebs, Amblyomma dissimile, Amblyomma ovale and Dermacentor nitens are present in both regions, humans have only been attacked by these species in the Neotropical Region. Even so, the Nearctic Region, despite its relatively limited ixodid fauna, is better represented by ticks that frequently attack humans than might be expected. By contrast, the Afrotropical Region, which contains the highest number of hard ticks found on humans, lags behind all other regions except the Australasian in terms of species found frequently or very frequently on people. For a given tick species, vectorial capacity varies between larvae, nymphs and adults, but, as noted above, morphological identification of the immature stages remains difficult. The recent rise of molecular taxonomy is likely to prove helpful in identifying tick immatures, but many larvae and nymphs collected from people will probably still not be determined to species, particularly when the collectors are not themselves trained in taxonomy, with the result that the number of identified species collected as immatures is liable to remain greatly underestimated. Immature specimens, even when often collected from people, will be identified only to genus or left undetermined, and we have excluded such specimens from this review. Of course, this is not to say that the identification of adult ticks is an easy task, and readers are cautioned that the records we have reported here are based on the literature, rather than examination of actual specimens, so the possibility exists that some identifica- tions are erroneous. Thus, Estrada-Peña et al. (2017b) found that misidentifications of Rhipicephalus species are common in Western Europe. Surely, this problem is not restricted to Western European workers, but it underscores the urgent need to refine current tick identification techniques. Comments and Conclusions 245

Table 6 summarizes our list of valid species (eight Prostriata and 22 Metastriata) that are treated in Chapter 2 as having been erroneously or dubiously regarded as human parasites. Where a species’ parasitological importance is unclear, further research should be conducted to ascertain whether humans are at risk and, if so, whether the species in question is a potential vector of pathogenic agents. Chapter 3 lists 107 invalid tick names that have been used historically, or in some cases currently, to describe tick species known to parasitize humans. This information should be of particular importance to nontaxonomists, who are often confused by the name changes that are an inevitable consequence of research in tick systematics.

Table 6 Ixodid species thought to have been wrongly, uncertainly or dubiously described as parasites of humans. The principal zoogeographic region is underlined. M: males, F: females, N: nymphs, A: adults but sex unknown, U: unknown. Tick species Zoogeographic distribution Tick stages Ixodes affinis Nearctic, Neotropical U I. hirsti Australasian U I. jellisoni Nearctic U I. laysanensis Central Pacific Islands U I. loricatus Neotropical A I. luciae Neotropical F I. simplex Afrotropical, Australasian, Oriental, F Palearctic I. trichosuri Australasian U Amblyomma auricularium Nearctic, Neotropical F A. clypeolatum Oriental U A. compressum Afrotropical A A. geayi Neotropical N A. helvolum Australasian, Oriental F, N A. macfarlandi Neotropical N A. pomposum Afrotropical U A. sylvaticum Afrotropical U A. usingeri Neotropical M, F Dermacentor halli Nearctic, Neotropical M D. limbooliati Oriental F D. taiwanensis Oriental U D. tamokensis Oriental M, F Haemaphysalis kashmirensis Oriental M, F H. muhsamae Afrotropical U H. palawanensis Oriental F H. warburtoni Palearctic M, F, N Hyalomma franchinii Afrotropical M, F H. impressum Afrotropical U Rhipicentor bicornis Afrotropical F Rhipicephalus dux Afrotropical F 246 Comments and Conclusions

Finally, in Table 7 we present a list of all the countries and territories from which tick species causing human parasitism have been reported, with those species frequently or very frequently found on people underlined. This table may be consid- ered a rough indicator of a country’s actual or potential tick threat to human health, but it almost certainly underestimates both the numbers and frequencies of tick species likely to parasitize people.

Table 7 Ixodid species found on humans, arranged by country/territory. Species found very frequently or frequently on people are underlined. Afghanistan: Dermacentor raskemensis. Algeria: Hyalomma dromedarii, H. impeltatum, H. lusitanicum, Rhipicephalus bursa, R. sanguineus s. l. Angola: Ixodes cavipalpus, Amblyomma sparsum Argentina: Ixodes pararicinus, Amblyomma aureolatum, A. brasiliense, A. coelebs, A. dubitatum, A. hadanii, A. incisum, A. neumanni, A. ovale, A. parvum, A. pseudoconcolor, A. pseudoparvum, A. sculptum, A. tigrinum, A. tonelliae, A. triste, Haemaphysalis juxtakochi, H. leporispalustris, Rhipicephalus microplus, R. sanguineus s. l. Australia: Ixodes australiensis, I. confusus, I. cornuatus, I. fecialis, I. holocyclus, I. kohlsi, I. myrmecobii, I. tasmani, Amblyomma limbatum, A. loculosum, A. moreliae, A. postoculatum, A. triguttatum, Bothriocroton auruginans, B. hydrosauri, Haemaphysalis bancrofti, H. humerosa, H. longicornis, H. novaeguineae, Rhipicephalus australis, R. sanguineus s. l. Austria: Ixodes ricinus Bangladesh: Demacentor auratus, Rhipicephalus microplus Belarus: Ixodes ricinus Belize: Ixodes boliviensis, Amblyomma coelebs, A. maculatum, A. mixtum Belgium: Ixodes hexagonus, I. ricinus, Rhipicephalus sanguineus s. l. Bolivia: Amblyomma parvum, Dermacentor nitens, Rhipicephalus microplus Bosnia and Herzegovina: Ixodes ricinus, Rhipicephalus sanguineus s. l. Brazil: Amblyomma aureolatum, A. brasiliense, A. cajennense, A. coelebs, A. dissimile, A. dubitatum, A. fuscum, A. incisum, A. latepunctatum, A. longirostre, A. naponense, A. oblongoguttatum, A. ovale, A. parkeri, A. parvum, A. romitii, A. rotundatum, A. scalpturatum, A. sculptum, A. tigrinum, Haemaphysalis juxtakochi, Rhipicephalus microplus, R. sanguineus s. l. Bulgaria: Ixodes ricinus, Haemaphysalis concinna, H. inermis, Hyalomma aegyptium, H. scupense, Rhipicephalus bursa, R. sanguineus s. l. Burkina Faso: Amblyomma tholloni, A. variegatum, Rhipicephalus guilhoni, R. sanguineus s. l., R. sulcatus Burundi: Rhipicephalus appendiculatus Cameroon: Ixodes rageaui, I. rasus, Amblyomma variegatum, Rhipicephalus complanatus, R. sanguineus s. l., R. ziemanni Canada: Ixodes angustus, I. cookei, I. dentatus, I. marxi, I. muris, I. pacificus, I. scapularis, I. soricis, I. spinipalpis, I. texanus, I. uriae, Dermacentor albipictus, D. andersoni, D. variabilis, Haemaphysalis chordeilis, H. leporispalustris, Rhipicephalus sanguineus s. l. Central African Republic: Amblyomma nuttalli, A. variegatum, Rhipicephalus sanguineus s. l., R. senegalensis, R. sulcatus (continued) Comments and Conclusions 247

Table 7 (continued) Chad: Rhipicephalus guilhoni, R. sanguineus s. l. Chile: Rhipicephalus sanguineus s. l. China: Ixodes acutitarsus, I. ovatus, I. persulcatus, I. sinensis, Amblyomma testudinarium, Demacentor auratus, D. niveus, D. nuttalli, D. silvarum, Haemaphysalis concinna, H. hystricis, H. japonica, H. longicornis, H. montgomeryi, H. nepalensis, H. qinghaiensis, Hyalomma asiaticum, H. dromedarii, Rhipicephalus bursa, R. haemaphysaloides, R. microplus, R. sanguineus s. l. Colombia: Amblyomma dissimile, A. mixtum, A. oblongoguttatum, A. ovale, Dermacentor imitans, D. nitens, Rhipicephalus microplus, R. sanguineus s. l. Congo: Rhipicephalus longus, R. sanguineus s. l. Costa Rica: Amblyomma coelebs, A. mixtum, A. ovale, A. pecarium, A. varium, Rhipicephalus microplus Croatia: Ixodes ricinus Cuba: Amblyomma mixtum Curação: Rhipicephalus sanguineus s. l. Cyprus: Haemaphysalis sulcata, Hyalomma anatolicum, H. excavatum Czech Republic: Ixodes ricinus, Haemaphysalis concinna, Dermacentor reticulatus Democratic Republic of Congo: Amblyomma tholloni, A. variegatum, Rhipicephalus appendiculatus, R. bequaerti, R. compositus, R. sanguineus s. l., R. supertritus Denmark: Ixodes ricinus, I. uriae Djibouti: Rhipicephalus sanguineus s. l. Ecuador: Amblyomma ovale Egypt: Hyalomma excavatum, H. impeltatum, Rhipicephalus sanguineus s. l. El Salvador: Rhipicephalus microplus Eritrea: Rhipicephalus sanguineus s. l. Ethiopia: Amblyomma falsomarmoreum, Haemaphysalis leachi, Hyalomma truncatum, Rhipicephalus pravus, R. pulchellus, R. sanguineus s. l. Fiji: Amblyomma breviscutatum, Finland: Ixodes persulcatus, I. ricinus France: Ixodes acuminatus, I. frontalis, I. hexagonus, I. ricinus, I. ventalloi, Dermacentor marginatus, D. reticulatus, Hyalomma lusitanicum, H. scupense, Rhipicephalus bursa, R. pusillus, R. sanguineus s. l., R. turanicus s. l. French Guiana: Amblyomma aureolatum, A. cajennense, A. longirostre, A. naponense, A. oblongoguttatum, A. ovale, A. tigrinum, Haemaphysalis juxtakochi, Rhipicephalus sanguineus s. l. Germany: Ixodes hexagonus, I. ricinus, Dermacentor marginatus, D. reticulatus, Haemaphysalis concinna, H. punctata, Rhipicephalus sanguineus s. l. Ghana: Ixodes rasus, I. vanidicus, Amblyomma tholloni, Dermacentor circumguttatus, Rhipicephalus complanatus, R. sanguineus s. l., R. ziemanni Great Britain: Ixodes canisuga, I. frontalis, I. hexagonus, I. ricinus, I. trianguliceps, I. uriae, I. ventalloi, Amblyomma hebraeum, Dermacentor reticulatus, Haemaphysalis punctata, Rhipicephalus sanguineus s. l. Greece: Ixodes gibbosus, I. ricinus, Dermacentor marginatus, Hyalomma anatolicum, H. excavatum, H. marginatum, H. rufipes, Rhipicephalus annulatus, R. bursa, R. sanguineus s. l., R. turanicus s. l. (continued) 248 Comments and Conclusions

Table 7 (continued) Guam: Amblyomma breviscutatum Guatemala: Amblyomma mixtum, A. parvum Guinea: Amblyomma variegatum Guinea-Bissau: Amblyomma variegatum, Rhipicephalus decoloratus Guyana: Amblyomma cajennense, A. oblongoguttatum, Rhipicephalus sanguineus s. l. Honduras: Amblyomma mixtum Hungary: Ixodes ricinus, Dermacentor marginatus, D. reticulatus Iceland: Ixodes ricinus, I. uriae India: Ixodes acutitarsus, I. kashmiricus, I. ovatus, I. petauristae, Amblyomma integrum, A. testudinarium, Demacentor auratus, Haemaphysalis aculeata, H. anomala, H. aponommoides, H. bispinosa, H. indoflava, H. montgomeryi, H. ramachandrai, H. spinigera, H. turturis, Hyalomma anatolicum, H. brevipunctatum, H. hussaini, H. isaaci, H. turanicum, Nosomma monstrosum, Rhipicephalus haemaphysaloides, R. sanguineus s. l., R. turanicus s. l. Indonesia: Ixodes spinicoxalis, Amblyomma babirussae, A. breviscutatum, A. geoemydae, A. testudinarium, Dermacentor auratus, D. steini, Haemaphysalis celebensis, H. hirsuta, H. hylobatis, H. mjoebergi, Rhipicephalus pilans, R. sanguineus s. l. Iran: Ixodes ricinus, Dermacentor raskemensis, Rhipicephalus bursa Irak: Hyalomma schulzei, Rhipicephalus sanguineus s. l. Ireland: Ixodes hexagonus, I. ricinus Israel: Ixodes redikorzevi, Haemaphysalis sulcata, Rhipicephalus bursa, R. sanguineus s. l. Italy: Ixodes acuminatus, I. festai, I. hexagonus, I. ricinus, I. ventalloi, Dermacentor marginatus, Haemaphysalis inermis, H. parva, H. punctata, H. sulcata, Hyalomma lusitanicum, H. marginatum, H. scupense, Rhipicephalus annulatus, R. bursa, R. pusillus, R. sanguineus s. l., R. turanicus s. l. Ivory Coast: Ixodes cumulatimpunctatus, I. muniensis, I. rasus, Amblyomma nuttalli, A. paulopunctatum, A. tholloni, A. variegatum, Dermacentor circumguttatus, Haemaphysalis parmata, Rhipicephalus aurantiacus, R. sanguineus s. l., R. senegalensis Jamaica: Amblyomma mixtum Japan: Ixodes acutitarsus, I. asanumai, I. columnae, I. monospinosus, I. nipponensis, I. ovatus, I. pavlovskyi, I. persulcatus, I. tanuki, I. turdus, Amblyomma geoemydae, A. testudinarium, Haemaphysalis campanulata, H. flava, H. hystricis, H. japonica, H. kitaokai, H. longicornis, H. megaspinosa, Rhipicephalus microplus, R. sanguineus s. l. Jordan: R. turanicus s. l. Kazakhstan: Ixodes pavlovskyi, I. persulcatus, Dermacentor marginatus, D. niveus, D. reticulatus, D. silvarum, Haemaphysalis erinacei, H. punctata, Hyalomma asiaticum, H. scupense, Rhipicephalus pumilio, R. schulzei Kenya: Ixodes schillingsi, Amblyomma hebraeum, A. personatum, A. tholloni, A. variegatum, Dermacentor rhinocerinus, Hyalomma albiparmatum, H. truncatum, Rhipicephalus appendiculatus, R. armatus, R. hurti, R. jeanneli, R. pravus, R. pulchellus, R. sanguineus s. l. Kosovo: Ixodes ricinus, Haemaphysalis punctata, Hyalomma marginatum Kyrgyzstan: Ixodes kazakstani, I. persulcatus, Dermacentor marginatus, Haemaphysalis caucasica, H. punctata, H. sulcata, Hyalomma anatolicum, H. marginatum, H. scupense, Rhipicephalus pumilio, R. sanguineus s. l., R. turanicus s. l. Laos: Haemaphysalis hystricis (continued) Comments and Conclusions 249

Table 7 (continued) Latvia: Ixodes ricinus Liberia: Rhipicephalus aurantiacus, R. complanatus Madagascar: Amblyomma variegatum, Haemaphysalis elongata, Rhipicephalus microplus Malaysia: Ixodes granulatus, Amblyomma cordiferum, A. geoemydae, A. testudinarium, Dermacentor compactus, D. steini, Haemaphysalis cornigera, H. nadchatrami, Rhipicephalus microplus, R. sanguineus s. l. Malawi: Amblyomma variegatum, Rhipicephalus appendiculatus, R. decoloratus, R. sanguineus s. l. Mali: Rhipicephalus sanguineus s. l. Mexico: Ixodes boliviensis, I. tancitarius, Amblyomma coelebs, A. dissimile, A. inornatum, A. maculatum, A. mixtum, A. ovale, A. parvum, A. tenellum, Dermacentor variabilis, Rhipicephalus microplus, R. sanguineus s. l. Moldova: Ixodes ricinus, Rhipicephalus rossicus Mongolia: Ixodes persulcatus, Hyalomma asiaticum, Rhipicephalus pumilio Morocco: Ixodes ricinus, Dermacentor marginatus, Hyalomma impeltatum, H. lusitanicum, Rhipicephalus sanguineus s. l., R. turanicus s. l. Mozambique: Amblyomma hebraeum, Hyalomma scupense, Rhipicephalus decoloratus, R. maculatus Myanmar: Ixodes acutitarsus, I. ovatus, Amblyomma testudinarium, Demacentor auratus, Haemaphysalis heinrichi, H. hystricis, Rhipicephalus haemaphysaloides Nepal: Ixodes acutitarsus, I. ovatus, Dermacentor auratus, Haemaphysalis anomala, H. aponommoides, H. birmaniae, H. montgomeryi, H. nepalensis, Nosomma monstrosum, Rhipicephalus haemaphysaloides, R. sanguineus s. l. Namibia: Hyalomma turanicum, Rhipicephalus gertrudae, R. pravus (as Rhipicephalus near pravus), R. simus, R. sulcatus Netherlands: Ixodes hexagonus, I. ricinus New Caledonia: Amblyomma loculosum, Rhipicephalus australis, R. sanguineus s. l. New Zealand: Haemaphysalis longicornis Nicaragua: Amblyomma mixtum Nigeria: Amblyomma variegatum, Hyalomma truncatum, Rhipicephalus muhsamae, R. sanguineus s. l. Norway: Ixodes ricinus, I. uriae Oman: Hyalomma dromedarii, Rhipicephalus sanguineus s. l. Pakistan: Dermacentor raskemensis, Hyalomma brevipunctatum Palau: Amblyomma loculosum Panama: Ixodes boliviensis, Amblyomma dissimile, A. mixtum, A. naponense, A. oblongoguttatum, A. ovale, A. parvum, A. pecarium, A. sabanerae, A. tapirellum, A. varium, Dermacentor imitans, D. latus, Haemaphysalis juxtakochi, Rhipicephalus sanguineus s. l. Papua New Guinea: Ixodes eichhorni, Haemaphysalis novaeguineae Papua New Guinea: Rhipicephalus sanguineus s. l. Paraguay: Amblyomma aureolatum, A. brasiliense, A. coelebs, A. incisum, A. ovale, A. tigrinum, A. tonelliae Peru: Rhipicephalus microplus, R. sanguineus s. l. (continued) 250 Comments and Conclusions

Table 7 (continued) Philippines: Amblyomma breviscutatum Poland: Ixodes hexagonus, I. persulcatus, I. ricinus, I. vespertilionis, Dermacentor reticulatus Portugal: Ixodes ricinus, Dermacentor marginatus, Hyalomma lusitanicum, H. marginatum, Rhipicephalus annulatus, Rhipicephalus bursa, R. pusillus, R. sanguineus s. l. Romania: Ixodes ricinus, Dermacentor marginatus, Haemaphysalis concinna, H. punctata, Hyalomma marginatum, Rhipicephalus bursa, R. rossicus, R. sanguineus s. l. Russia: Ixodes angustus, I. apronophorus, I. crenulatus, I. laguri, I. nipponensis, I. pavlovskyi, I. persulcatus, I. ricinus, I. uriae, Dermacentor marginatus, D. niveus, D. nuttalli, D. reticulatus, D. silvarum, Haemaphysalis concinna, H. inermis, H. japonica, H. longicornis, H. parva, H. punctata, H. sulcata, Hyalomma asiaticum, H. marginatum, H. scupense, Rhipicephalus pumilio, R. rossicus, R. schulzei, R. sanguineus s. l., R. turanicus s. l. Rwanda: Ixodes cumulatimpunctatus Saudi Arabia: Hyalomma excavatum, H. rufipes, Rhipicephalus sanguineus s. l., R. turanicus s. l. Senegal: Amblyomma variegatum, Hyalomma truncatum, Rhipicephalus sanguineus s. l., R. sulcatus Serbia: Ixodes ricinus, Haemaphysalis concinna Seychelles: Amblyomma loculosum Sierra Leone: Rhipicephalus lunulatus, R. sanguineus s. l., R. senegalensis Slovakia: Dermacentor marginatus, D. reticulatus, Haemaphysalis inermis, H. punctata Somalia: Rhipicephalus humeralis, R. pravus, R. pulchellus South Africa: Ixodes pilosus, I. rubicundus, Amblyomma hebraeum, A. marmoreum, Haemaphysalis elliptica, H. parmata, H. silacea, Hyalomma glabrum, H. rufipes, H. truncatum, Rhipicephalus appendiculatus, R. decoloratus, R. follis, R. gertrudae, R. glabroscutatus, R. maculatus, R. microplus, R. muehlensi, R. sanguineus s. l., R. simus, R. warburtoni, R. zambeziensis South Korea: Ixodes granulatus, I. nipponensis, I. persulcatus, Amblyomma testudinarium, Haemaphysalis flava, H. japonica, H. longicornis South Sudan: Amblyomma lepidum, A. variegatum, Rhipicephalus appendiculatus, R. muhsamae, R. pravus, R. sanguineus s. l. Spain: Ixodes ricinus, Dermacentor marginatus, D. reticulatus, Haemaphysalis punctata, Hyalomma marginatum, Rhipicephalus annulatus, R. bursa, R. pusillus, R. sanguineus s. l., R. turanicus s. l. Sri Lanka: Ixodes petauristae, Amblyomma integrum, A. testudinarium, Demacentor auratus, Haemaphysalis aculeata, H. bispinosa, H. cuspidata, H. intermedia, H. spinigera, H. turturis, Hyalomma brevipunctatum, H. isaaci, Rhipicephalus haemaphysaloides, R. microplus, R. sanguineus s. l. Sudan: Amblyomma lepidum, Hyalomma rufipes, Rhipicephalus sanguineus s. l. Suriname: Amblyomma cajennense, A. oblongoguttatum, A. ovale, A. pacae Sweden: Ixodes ricinus, Haemaphysalis punctata Switzerland: Ixodes ricinus, Dermacentor marginatus, Haemaphysalis punctata, Rhipicephalus sanguineus s. l. (continued) Comments and Conclusions 251

Table 7 (continued) Taiwan: Ixodes acutitarsus, Amblyomma testudinarium, Haemaphysalis hystricis, Rhipicephalus haemaphysaloides, R. sanguineus s. l. Tajikistan: Dermacentor marginatus, Haemaphysalis sulcata, Hyalomma anatolicum, H. asiaticum, H. scupense, H. turanicum, Rhipicephalus annulatus Tanzania: Ixodes cumulatimpunctatus, Amblyomma tholloni, A. variegatum, Rhipicephalus evertsi, R. kochi, R. planus, R. sanguineus s. l., R. sulcatus Thailand: Ixodes acutitarsus, I. granulatus, Amblyomma javanense, A. testudinarium, Demacentor auratus, D. steini, Haemaphysalis anomala, H. bispinosa, H. darjeeling, H. doenitzi, H. hystricis, H. koningsbergeri, H. lagrangei, H. nadchatrami, H. obesa, H. papuana, H. semermis, H. shimoga, H. wellingtoni, Rhipicephalus haemaphysaloides, R. sanguineus s. l. Trinidad and Tobago: Amblyomma calcaratum, A. mixtum Turkey: Ixodes acuminatus, I. gibbosus, I. hexagonus, I. laguri, I. redikorzevi, I. ricinus, Dermacentor marginatus, D. niveus, D. reticulatus, Haemaphysalis concinna, H. erinacei, H. inermis, H. parva, H. punctata, H. sulcata, Hyalomma aegyptium, H. anatolicum, H. excavatum, H. impeltatum, H. marginatum, H. rufipes, H. scupense, H. turanicum, Rhipicephalus annulatus, R. bursa, R. sanguineus s. l., R. turanicus s. l. Turkmenistan: Rhipicephalus turanicus s. l. Uganda: Amblyomma cohaerens, A. tholloni, Dermacentor circumguttatus, Haemaphysalis paraleachi, Rhipicephalus appendiculatus, R. longus, R. lunulatus, R. praetextatus, R. sanguineus s. l. Ukraine: Ixodes redikorzevi, I. ricinus, Dermacentor marginatus, Haemaphysalis inermis, H. punctata, Hyalomma marginatum, H. scupense, R. rossicus, R. sanguineus s. l. Uruguay: Amblyomma dubitatum, A. tigrinum, A. triste, Haemaphysalis juxtakochi, Rhipicephalus sanguineus s. l. USA: Ixodes angustus, I. baergi, I. banksi, I. brunneus, I. cookei, I. dentatus, I. kingi, I. marxi, I. muris, I. pacificus, I. rugosus, I. scapularis, I. sculptus, I. spinipalpis, I. texanus, I. uriae, I. woodi, Amblyomma americanum, A. inornatum, A. latum, A. maculatum, A. mixtum, A. tenellum, A. triste, A. tuberculatum, Dermacentor albipictus, D. andersoni, D. hunteri, D. occidentalis, D. parumapertus, D. variabilis, Haemaphysalis chordeilis, H. leporispalustris, Rhipicephalus annulatus, R. sanguineus s. l. Uzbekistan: Hyalomma anatolicum Vanuatu: Amblyomma breviscutatum Venezuela: Amblyomma dissimile, A. longirostre, A. oblongoguttatum, A. ovale, A. triste, Dermacentor nitens, Haemaphysalis juxtakochi, Rhipicephalus microplus, R. sanguineus s. l. Vietnam: Ixodes granulatus, Amblyomma testudinarium, Demacentor auratus, D. bellulus, D. compactus, Haemaphysalis colasbelcouri, H. hystricis, H. lagrangei, H. roubaudi, H. shimoga, Rhipicephalus microplus, R. sanguineus s. l. Yemen: Amblyomma variegatum, Hyalomma rufipes, Rhipicephalus sanguineus s. l. Zambia: Ixodes cavipalpus Zimbabwe: Ixodes cumulatimpunctatus, Amblyomma hebraeum, A. nuttalli, Rhipicephalus sanguineus s. l. (continued) 252 Comments and Conclusions

Table 7 (continued) UNDETERMINED COUNTRIES Undetermined African countries: Ixodes cavipalpus, Amblyomma gemma, A. hebraeum, A. lepidum, A. marmoreum, A. tholloni, A. variegatum, Hyalomma truncatum, Rhipicephalus appendiculatus, R. armatus, R. carnivoralis, R. complanatus, R. compositus, R. decoloratus, R. distinctus, R. evertsi, R. fulvus, R. humeralis, R. hurti, R. jeanneli, R. longus, R. lunulatus, R. maculatus, R. muhsamae, R. planus, R. praetextatus, R. pravus, R. pulchellus, R. sanguineus s. l., R. senegalensis, R. simus, R. sulcatus, R. supertritus, R. turanicus s. l., R. ziemanni, R. zumpti Undetermined American countries: Amblyomma ovale, A. tapirellum, A. variegatum Undetermined Asian countries: Dermacentor auratus, D. niveus, Haemaphysalis hylobatis, H. obesa, Hyalomma hussaini, H. isaaci, Rhipicephalus haemaphysaloides, R. pilans, R. pumilio, R. schulzei Undetermined northwest European countries: Ixodes acuminatus, I. hexagonus, I. ricinus, I. trianguliceps, I. uriae, I. ventalloi, Dermacentor marginatus, D. reticulatus, Haemaphysalis concinna, H. inermis, H. punctata, Rhipicephalus sanguineus s. l. Undetermined South American countries: Amblyomma aureolatum Undetermined former Yugoslavian countries: Ixodes ricinus, Haemaphysalis punctata, H. sulcata, H. scupense, Rhipicephalus bursa, R. sanguineus s. l. Undetermined countries: Amblyomma variegatum, Hyalomma asiaticum, H. dromedarii, Rhipicephalus bursa, R. rossicus, R. sanguineus s. l., R. turanicus s. l. References

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