Perception of Ticks and Tick-Borne Diseases Worldwide: Comparison
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Please note this is a comparison between Version 2 by Lindsay Dong and Version 1 by Jose De La Fuente.

In humans, an obvious reduction of the impact of ticks and tick-borne diseases (TBDs) could be managed by informing the population on the risks associated with ticks and TBD, involving frequent public news media and advertisements, as currently carried out in northern countries of Europe, which are measuring their impact and adaptation [254]. Although the U.S. Department of Agriculture (USDA), the National Institutes of Health (NIH), and the European Centre for Disease Prevention and Control provide online free access to information about TBD, gaps are obvious in both the transmitted information and the ability of citizens to understand the information. The same applies to ticks feeding on pets, that have an extraordinarily high contact with humans.

  • tick
  • tick-borne diseases
  • environment
  • surveillance

1. Introduction

Ticks and tick-borne diseases (TBDs) are a growing burden worldwide with (re)emerging diseases affecting human and animal health (e.g., recent references [1,2,3,4,5,6,7][1][2][3][4][5][6][7]). Factors behind this increase in cases, detection of new pathogens, or new epidemics in areas previously free of a pathogen are varied and sometimes of a local nature (i.e., [8,9][8][9]). For example, the trends of climate have been mentioned as the source of spread of some species of ticks [10,11][10][11]; the availability of meta-genomics improved the detection of previously unknown tick-borne viruses [12,13][12][13]. The changes in the landscape derived from human actions (e.g., changes in culture patterns, abandonment of culture areas, deforestation in some zones of South America) have been indicated as the main reason for epidemics of tick-borne pathogens in both animals and humans [14].
Hominids evolved in interaction with ticks and TBD as supported by fossil tick amber inclusions dated at ca. 100 Mya (Cretaceous), estimated origin of Ixodida at ca. 350 Mya, and the presence of TBPs in fossil ticks [15,16,17,18,19][15][16][17][18][19]. However, while some non-human primates have specific species of ticks, the same does not hold for species of ticks parasitizing the genus Homo, and Homo sapiens lacks its “own” species of ticks. All the species of ticks affecting humans (compiled by [20]) are either generalist species or the result of an accidental parasitism of ticks with varied specificity (ruminants, carnivores). On the other hand, the pathogens carried by these species affecting non-human primates have been seldom investigated, like the Kyasanur forest virus and Haemaphysalis bispinosa.
The conclusion is that both adequate passive and active surveillance, according to the logistic issues or other circumstances [21] regarding ticks affecting humans, and an active study of ticks affecting livestock and pets, aiming to improve both the health of the animals and the economic outcome, are necessary. However, despite advances in the surveillance, epidemiology, identification/diagnostics, and preventive/control interventions for ticks and TBD, major challenges are faced due to global expansion and increased incidence of TBD. One of these challenges is the difference that may exist in the perception of ticks and TBD worldwide. This perception is impossible to capture without the view of experts in different countries and regions.

2. Contributions from Different Countries and Regions Worldwide

2.1. United States of America

Ticks and tick-borne disease constitute a growing burden in the United States (US) in both residential urban and land environments [10,22,23][10][22][23]. According to a recent publication by Eisen (2022) [10], 36 ixodid (most recorded, Ixodes scapularis, Amblyomma americanum, Dermacentor variabilis, Ixodes pacificus, and Dermacentor andersoni) and 13 argasid species (most recorded, Otobius megnini and Ornithodoros coriaceus) have been associated with human infestations in the US. Other tick species recorded in humans (>250 records) included Ixodes cookei, Dermacentor occidentalis, Rhipicephalus sanguineus s.l., Dermacentor albipictus, and Amblyomma maculatum [10]. The most recorded tick species in humans representing 67% of all ixodid tick records is I. scapularis, vector of pathogens and associated diseases, Borrelia burgdorferi senso stricto and Borrelia mayonii (Lyme disease), Borrelia miyamotoi (hard tick-borne relapsing fever), Anaplasma phagocytophilum (human granulocytic anaplasmosis), Ehrlichia muris eauclairensis (ehrlichiosis), Babesia microti (babesiosis), and Powassan virus (Powassan encephalitis) [10]. Factors such as climate change drive the expanding geographical range in the US of tick species such as A. americanum and I. scapularis and thus the incidence of TBDs such as anaplasmosis, babesiosis, Lyme disease, ehrlichiosis, and arboviral diseases [9,10,21][9][10][21]. The increased incidence of alpha-gal syndrome (AGS) has also been associated with expanding A. americanum [10] and is underdiagnosed [25,26][24][25]. AGS is an emerging multisymptomatic allergic disease mediated by IgE-type antibody response to galactose-alpha-1,3-galactose (alpha-gal) and associated with tick bites and consumption of mammalian meat and derived products containing alpha-gal [25,26,27][24][25][26]. Personal protection measures to prevent human contact with ticks and thus reduce the risk of tick bites are highly recommended to be used consistently [28,29][27][28]. According to Eisen (2022) [29][28], protection measures include “use of repellents, wearing untreated or permethrin-treated protective clothing, and conducting tick checks after coming inside, aided by removing outdoor clothing articles and running them in a dryer on high heat (to kill undetected ticks) and taking a shower/bath (to aid in detecting ticks on the skin)”. Other protection measures to consider include landscaping, vegetation management, tick host fencing, use of four-poster tick control deer feeders to apply acaracide to white-tailed deer, deer herd reduction, implementation of i-tree canopy vegetation cover subtype classification to predict peri-domestic tick presence, pet tick control, and interventions to kill host-seeking ticks or ticks infesting rodents [11,22,30,31,32,33,34][11][22][29][30][31][32][33]. Informing the population on the risks associated with ticks and TBD and targeted education for the implementation of protection measures through social media and advertisements is important to reduce the incidence of TBD [28,35][27][34]. Although the U.S. Department of Agriculture (USDA, Washington, DC, USA) and the National Institutes of Health (NIH, Bethesda, MD, USA) provide online free access information about TBD, gaps in population knowledge and differences in the attitudes and motivation such as forgetfulness, safety concerns, and lack of awareness affect the implementation of protection measures [36,37,38][35][36][37].  Gaps in the diagnosis, prognosis, treatment, and prevention of TBDs are a limitation for the reduction of the incidence and severity associated with these diseases [44][38]. Laboratory diagnostic methods are not well implemented and not effective for diagnosis during the acute illness stage when timely treatment is needed, while nucleic acid amplification tests are most effective [45][39]. 

2.2. Mexico

Ticks and tick-borne diseases are a significant concern in Mexico. The two most common tick species affecting domestic animals in the country are the hard ticks Rhipicephalus microplus and R. sanguineus. R. microplus is found in over 60% of Mexico, while R. sanguineus is more widely distributed [46,47][40][41]. Other tick species such as Amblyomma spp., Dermacentor spp., and Ixodes spp. can also be found. Otobius megnini has been frequently found parasitizing cattle and less regularly found on dogs and horses [48,49][42][43]. Babesiosis and anaplasmosis are the most prevalent TBDs in cattle, with prevalence rates ranging from 2% to 94% and >50%, respectively [50,51][44][45]. These diseases constantly threaten livestock and limit beef cattle’s genetic improvement due to the high morbidity and mortality of high-value animals introduced to tick-infested areas [50][44]. Equine babesiosis and theileriosis are prevalent in horses, complicating their movement and transportation for sport, competition, and as companion equids [51][45]. Rickettsiosis by R. rickettsii is the most important TBD in humans in northwestern Mexico, with mortality rates of 30–40% [54][46]. In addition, infection with B. burgdorferi has been confirmed in over 100 cases [55][47]. Although evidence of human babesiosis and anaplasmosis has been documented for a long time, only recently have B. microti and A. phagocytophilum been molecularly identified [56,57][48][49]. However, the tick vector remains to be determined. Cattle producers, especially those in northern Mexico and the Gulf of Mexico, know the importance of R. microplus due to the national campaign against this tick. However, a wide gap in education and training for other tick species still needs to be addressed. Therefore, misunderstanding and need for knowledge on the role of ticks as vectors of pathogens of zoonosis concern in both rural and suburban areas exist. Outbreaks of TBD during the season with the highest abundance of brown dog ticks in the northwest of the country are an example of the lack of information on preventing tick infestations and controlling TBD. The prevention and control of ticks and the diseases they transmit to animals and humans require a research agenda that considers tick biology, integrated tick control, and science-based use of acaricides alone and combined with anti-tick vaccines together with management of wild animal translocation, tick surveillance, identification of tick carriers and reservoirs, standardization of diagnosis methods, and molecular identification of pathogens [58][50].

2.3. Central America

Central America contains an approximate area of 522,000 km2 integrating a wide biological diversity, which includes a rich fauna of ticks, with about 80 species reported to date (Supplementary Dataset S3). Of this diversity of ticks, Amblyomma mixtum, Amblyomma ovale, Dermacentor nitens, R. microplus, Rhipicephalus sanguineus s.l., Alectorobius puertoricensis, and Alectorobius talaje have been reported as relevant to human and animal health due to their role as vectors of pathogens causing rickettsiosis, ehrlichiosis, anaplasmosis, relapsing fever, and babesiosis, as well as causing paralysis and severe allergies. Central America has a long-standing history in relation to cases of human and animal parasitism, with the first records of effects on humans being recorded in the 19th century in Guatemala (Ornithodoros talaje and Amblyomma sabanerae). At the beginning of the 20th century, the first reports of clinical cases of tick-borne pathogens in humans were reported in Panama (Rickettsia rickettsii relapsing fever and spotted fever) and Costa Rica (R. rickettsii spotted fever). In fact, rickettsiosis is the most important group of diseases reported in Central America, since there are confirmed fatal cases in these two countries, in addition to reports of severe rickettsiosis in acute patients from Guatemala, Honduras, and Nicaragua. Other microorganisms detected in ticks from Central America include the genera Ehrlichia (E. canis, E. cf. chaffeensis), Anaplasma (A. marginale, A. phagocitophylum, A. platys), Borrelia (B. puertoricensis, Borrelia burgdorferi group), and hemoparasites like Babesia (B. odocoilei, B. vogeli) and Hepatozoon (H. canis, Hepatozoon spp). There is also serologic evidence of ehrlichiosis, anaplasmosis, and babesiosis in domestic animals; a probable case of canine ehrlichiosis in a boy from Panama and serology compatible with ehrlichiosis in human blood from Costa Rica. 

2.4. Brazil

Brazil is a vast and ecologically diverse country with several distinct biomes that include tropical forests (Amazon and Atlantic rainforest), savannah (Cerrado), grasslands (Pampa), semi-arid (Caatinga), and the world’s largest tropical wetland (Pantanal). These biomes are characterized by their unique climate, vegetation, and wildlife. However, huge areas within each biome were transformed into anthropogenic landscapes to become part of the Global Human Ecosystem. This ecological mosaic has shaped the current Brazilian tick fauna and associated microbiota, but general knowledge about most tick species is lacking, and epidemiological data about transmitted pathogens are also scarce. Indeed, knowledge of tick-borne diseases is primarily related to those agents with a major impact on human welfare. The tick fauna of the country is by now composed of 78 species, 53 Ixodidae and 25 Argasidae. Amblyomma remains as the richest, with 34 valid species [59][51]. The original tick fauna was modified by the introduction of exotic species, outstandingly R. microplus and two species of the Rhipicephalus sanguineus complex [60,61,62][52][53][54]. Undoubtedly, the cattle tick R. microplus is a species that raises an important level of apprehension. It is the species most associated with economic losses throughout a country that had a commercial herd estimated at 224.6 million heads in 2022 [63][55]. In the last broad assessment of the negative impact of the cattle tick, an annual loss of USD 3.24 billion was estimated [64][56]. This loss includes the negative impact of infections caused by the major cattle tick-borne pathogens Babesia spp. and Anaplasma marginale and the disease commonly known as “Bovine parasitic sadness” [65][57].  The main horse ticks in the country are Amblyomma sculptum and Dermacentor nitens (named previously Amblyomma cajennense and Anocentor nitens) [68][58]. Among tick-borne diseases, equine piroplasmosis caused by Babesia caballi and Theileria equi infection is enzootic in Brazil [69,70][59][60]. Several tick species are supposed to transmit these pathogens but D. nitens is considered the sole vector of B. caballi, while T. equi is transmitted by R. microplus and possibly by A. sculptum [70,71][60][61].  In relation to dogs, the anthropogenic landscapes throughout Brazil are widely colonized by ticks of the R. sanguineus complex with a wide distribution in anthropized areas of the country [61][53]. These species, particularly the tropical lineage (recently considered Rhipicephalus linnaei [73][62]), are vectors of Ehrliquia canis and Babesia vogeli, the agents, respectively, of canine monocytic ehrlichiosis and canine piroplasmosis, collectively known by pet owners as the “tick disease” (in Brazilian Portuguese, “doença do carrapato”). Both E. canis and B. vogeli have been widely reported in dogs from Brazil [74,75][63][64]. These tick species have been found infected with R. rickettsii [76][65], nonetheless human rickettsiosis caused by infected tick bite remains elusive. On the other hand, Amblyomma aureolatum, a species restricted to the Atlantic rainforest and the Pampa biome in the south of the country [77[66][67],78], is the natural vector of Rangelia vitalii, the etiologic agent of canine rangeliosis, the most severe canine piroplasmosis of the western hemisphere [79][68].  Dogs are also involved in the epidemiology of human tick-borne rickettsiosis and should be considered a target species for tick control. Only two tick-borne Rickettsia species have been proven to cause human disease in Brazil, R. rickettsii, causing a frequently lethal disease, and Rickettsia parkeri strain Atlantic rainforest, responsible for milder non-lethal rickettsiosis [84][69]. Circumstantial evidence indicates that there is in the country a third and mild rickettsiosis caused by Rickettsia parkeri stricto sensu [85][70]. Wild carnivores are hosts for the adult ticks of Amblyomma aureolatum, Amblyomma ovale, and Amblyomma tigrinum and domestic dogs are common alternative hosts [86][71]. These tick species have been shown to be infected with, respectively, R. rickettsii, Rickettsia parkeri Atlantic rainforest strain, and Rickettsia parkeri sensu stricto and emerging knowledge indicates that dogs may bridge the infected ticks to human households [85,87,88][70][72][73]. Whereas Rickettsia-infected A. ovale ticks have a wide distribution within the country, infected A. tigrinum were detected only in the southern region [85][70].  Rickettsia rickettsii infection is the major human tick-borne disease in Brazil, the “febre maculosa Brasileira” (Brazilian spotted fever). Although the disease has a low prevalence and is overwhelmingly restricted to specific areas, it has gained significant attention and apprehension in society due to its high lethality. In fact, timely and correct antibiotic treatment is curative. Unfortunately, early diagnosis is not easy and relies on epidemiological data (febrile individuals bitten by ticks in endemic areas) since laboratory diagnosis is typically confirmatory after the recovery or death of those who are ill [90][74].  A controversial tick-borne disease in Brazil is Lyme borreliosis. Lyme-like disease has been diagnosed since 1992 [95,96][75][76]. The disease is routinely diagnosed based on clinical and serological data and records of suspected cases included in the Brazilian Ministry of Health database [96][76]. There are also occasional molecular identification reports of bacteria from the Borrelia burgdorferi sensu lato complex [97][77]. However, Borrelia burgdorferi has not yet been isolated either from humans or ticks [93,94][78][79] and the ecological background to sustain its epidemiology within the country is weak.  Viruses are also significant tick-borne agents, and tick-associated viruses have already been documented in Brazil [103,104][80][81]. However, their role as pathogens remains uncertain. Indeed, there are numerous molecular studies reporting other potential tick-borne pathogens in ticks collected from domestic animals, wild animals, and the environment in Brazil.

2.5. Europe

Ticks are an important part of the parasitic burden affecting livestock in Europe, as well as a growing issue regarding human health because of the transmission of pathogens. In Europe, prominent species of ticks affecting domestic animals (with even 6–7 generations per year, like R. microplus in many parts of the world) are absent, but reported species also represent an important burden in animal husbandry. The panorama is a wide repertoire of species, most of them affecting livestock, that colonize areas with very different environmental conditions therefore resulting in a “mosaic” of distribution [108,109,110[82][83][84][85][86],111,112], with different seasonal activity periods, ability to transmit different types of pathogens, and capacity to spread throughout the wild fauna of a region. Most of these ticks have generalist feeding habits, affecting notably domestic ruminants and horses under extensive management. They constitute a large burden affecting the production of meat or milk, debilitating the animals and/or increasing abortions, favoring poor health conditions, and promoting the development of secondary diseases caused by opportunistic bacteria [111][85]. An additional issue is the use of acaricides against ticks, which contribute to contamination by these toxic products and the increase in the costs of management of the animals. It is important to note that most (if not all) species of ticks affecting livestock are shared with wild ungulates. Therefore, due to the co-existence of wildlife and livestock in large European regions, efforts to control or eradicate ticks are challenging. As in other regions, ticks prevail in nature through cycles of infestation affecting either domestic or wild ungulates as adults, with the immatures feeding commonly on many species of birds and rodents [112][86]. Europe can be roughly divided, according to latitude, into three regions, Mediterranean, Central, and Northern regions. The Mediterranean region is populated by species of ticks with an obvious seasonality because of the seasonal nature of the weather in the region. The most important species belong to the genera Rhipicephalus and Hyalomma. Due to the vegetal characteristics of the region, sheep and goats are the main livestock present in the area (high humidity deficit, high temperature), making these species the main vectors for several species of protozoans, like Babesia spp. and Theileria spp., or bacteria like Anaplasma ovis or Rickettsia spp. Central Europe, including the British Isles and southern parts of Scandinavia and Finland, is the major area of distribution of the prominent species I. ricinus and H. punctata. These two species tend to concentrate on ruminants and are the main vectors of several species of protozoans of the genus Babesia and the bacterium Anaplasma phagocytophilum. Both pathogens are responsible for a wide array of clinical presentations, from the chronic one, with a course of abortions and serious weight loss, to the acute cases, in which death may be fast, even in 72 h. No efforts to determine the economic losses produced by these protozoans or bacteria have been addressed. Most, if not all, species affecting livestock and pets in Europe may bite humans with a different pressure according to the climate gradient associated with the territory and with pathogen transmission. The pathogens carried by ticks are supported by populations of wild vertebrates, that have a different prominence according to the composition of the community of vertebrates [117][87]. Other than pets, for which harmonized guidelines about tick control exist and owners commonly follow recommendations by veterinarians, there are no agreed protocols for tick control in Europe. Control (or attempt at eradication) of ticks depends on the perception of the owners, the recommendation of field veterinarians, and the availability of adequate acaricides, which may have different regulations depending on the country. The lack of harmonized protocols prevents the necessary joint effort to eradicate ticks. 

2.6. Egypt

In Egypt, animal trade, climate, and anthropogenic factors contribute to the spread of tick species and TBD. The spillover of various tick-borne pathogens is likely to occur from sub-Saharan Africa and other Mediterranean basin countries. The development of acaricide resistance further exacerbates the widespread presence of ticks and TBD, posing a significant economic challenge in Egypt [119,120][88][89]. To date, eight tick species from the family Argasidae and forty-four species from the family Ixodidae have been reported, with Hyalomma sp. and Rhipicephalus sp. being the most common. Enhancing the understanding of ticks and their associated diseases among various societal sectors, as well as the general population, is crucial for implementing effective control measures. In Egypt, ticks and TBD have predominantly been viewed through the lens of agricultural production rather than human health. Farmers’ perception is limited due to the widespread use of acaricides to eradicate ticks infesting animals, leading to the development of acaricide resistance [121,122][90][91]. Considering the close interaction between humans, animals, and tick vectors, a multidisciplinary approach linking human, animal, and environmental health within a “One Health” framework is essential. Systematic and comprehensive surveillance studies investigating ticks and TBD in defined areas are needed. Collaborative efforts between Egypt and Europe, combining fieldwork, research capacity, and funding, could lead to a better understanding of the epidemiological landscape of ticks and TBD. 

2.7. Uganda

In Uganda, ticks including Rhipicephalus appendiculatus, Rhipicephalus decoloratus, Amblyomma variegatum, and Rhipicephalus evertsi evertsi are the most economically important ticks that parasitize cattle and transmit deadly disease pathogens. The key tick-borne disease pathogens are Theileria parva, Babesia bovis, B. bigemina, A. marginale, and Ehrlichia ruminantium whose infections result in high morbidity and mortality if naïve cattle become infected. These diseases and the tick vectors cause annual losses of USD 1.1 billion, thus affecting cattle-keeping communities in poverty. However, the current tick control approaches mainly depend on the use of acaricides applied at a frequency of one to three times a week depending on the extent of the tick burden. The increasing frequency of application is critically indicative of acaricide-resistant tick genotypes [123][92]. The specific deleterious effects of tick infestations are bites which damage the hides in animals with high tick loads, blood loss and thus anemia, allergy due to toxins in tick saliva, chronic stress, and continuous irritation affecting animal welfare, leading to immuno-depression and loss of energy [124][93].  Generally, more than 70% of Uganda’s population depends on agriculture for their livelihood, and the animal industry accounts for 17% of the national gross domestic product. Cattle farmers perceive ticks and tick-borne diseases as a big limitation to commercial livestock farming given the fact that transmitted pathogens limit the breeding of high-yielding cattle. The overdependence on acaricide for tick control is no longer viable, thus demanding the development of novel control strategies.

2.8. Nigeria

In Nigeria, the study of ticks and tick-borne diseases has a history spanning several decades [125,126,127,128][94][95][96][97]. Although there was a period without tick research, there has been a recent resurgence of interest in investigating various tick-borne pathogens in Nigeria [129,130,131,132,133,134,135][98][99][100][101][102][103][104] and in collaboration with five other African countries [136,137][105][106]. Various studies have reported on the distribution of ticks on different animal hosts across the country, including wild game animals and cattle that enter through trans-border routes [130,131,132,138,139,140,141,142,143,144][99][100][101][107][108][109][110][111][112][113]. Regarding tick-borne diseases, babesiosis, anaplasmosis, theileriosis, and ehrlichiosis have received the most attention in documented cases [145,146][114][115]. However, conditions like CCHF and African tick-bite fever have been underreported, possibly indicating a gap in surveillance and reporting mechanisms for these particular diseases. Also, hundreds of ticks have been gathered from a snake kept in a zoo. All the ticks harvested were Amblyomma latum of both sexes and at different stages of development. Hepatozoon phythonis was identified by thin blood smear from the same snake, while Amblyomma tholloni was found on an elephant calf that was to be kept in a private zoo in the state of Edo, Nigeria.

2.9. India

In India, 109 tick species are reported to infest animals [150][116]. The tick index or tick burden in cattle was reported to be 0.922 to 1.0 [151,152][117][118] and the species diversity was high in rural parts in comparison to urban areas. As per a recent estimate, the TBD in animals is causing an economic loss of USD 787.63 million/year. Besides animals, several tick species such as Amblyomma integrum, Haemaphysalis spinigera, Dermacentor auratus, Hyalomma isaaci, Rhipicephalus haemaphysaloides, R. sanguineus s.l., and Otobius megnini are reportedly infesting human beings [153,154][119][120]. The use of acaricides by swabbing, dipping, spraying, pour-on, spot-on, and injection is the sole approach adopted for tick control. Awareness on environmental tick control or off-the-host tick control is lacking. No commercial anti-tick vaccines against the major cattle tick R. microplus/H. anatolicum are available which could be attributed to the diversity of targeted antigen sequences across the tick isolates in India [155][121]. The TBDs affecting humans are Kyasanur forest disease (KFD), CCHF, Ganjam virus (GANV), Bhanja virus (BHAV), Lyme disease, Q fever, rickettsial infections (Rickettsia conorii and R. rickettsii), and babesiosis (Babesia microti) [158][122]. Meanwhile, animals suffer from theileriosis, babesiosis, anaplasmosis, ehrlichiosis, hepatozoonosis, and lumpy skin disease virus. There are only two licensed vaccines available against TBD in India, viz., tropical bovine theileriosis (Rakhsavac-T) and KFD for humans. Chemotherapy is the only option being practiced, controlling major TBD infections. In rural communities, tick infestation is considered as one of the many problems animals have to suffer perennially and that is managed by washing animals, by rubbing them with dry fodder, or by the application of available acaricides at the local market when infestation is visible. In the organized sector, highly tick-susceptible cross-bred animals are maintained for higher production. The problem of ticks is regularly treated by the use of acaricides but without a strict adherence to the recommendation of the manufacturers. Although farmers are well aware of the high cost involved in the treatment of TBD, due to limited knowledge of the methods for tick control, resource-poor farmers face severe economic distress. 

2.10. Nepal

In the Nepalese context, there are inadequate studies on hard ticks and hard tick-borne diseases (HTBDs). In the hills and plains of western and central Nepal, the abundant cattle ticks are Rhipicephalus (Boophilus) microplus, Haemophysalis spp., Ixodes spp., and Amblyomma spp. [159,160][123][124]. Six species of Haemophysalis, five species of Rhipicephalus, and one species each of Amblyomma and Ixodes were reported from goats of Chitwan District [161][125]. Interestingly, two species of hard ticks (Amblyomma grevaisi and Amblyomma varanense) were identified in snakes of Nepal, and Amblyomma grevaisi was detected almost 100 years ago [162][126]. Similar to hard tick studies, scanty research has been carried out in Nepal on characterizing the HTBDs. In a serological study in Banke and Surkhet Districts [159][123], a 6.4% infection rate of TBDs was reported in cattle with Anaplasma marginale (5.8%) followed by Babesia bovis (0.6%). Rickettsia honei was reported in one human patient infested with ticks [166][127]. Lyme disease (caused by the tick-transmitted spirochaete Borrelia burgdorferi) was reported for the first time in 2018 in Kaski District [167][128] and a subsequent case was reported in a patient from Gulmi District [168][129]. Canines transmit vector-borne diseases at the wild–domestic interface. Particularly, infections in stray dogs are alarming in Kathmandu Valley, where 81.43% of the stray dogs are infected by at least one vector-borne pathogen (Anaplasma platys (31%), Babeisa vogeli (13%), Babesia gibsoni (23%)) and 41.43% are co-infected with more than one vector-borne disease [169][130]. 

2.11. Indonesia

Geographically, Indonesia is an archipelagic country on the equator, known as having almost the most biodiversity in the world, second only to the Brazilian Amazon. The islands in eastern Indonesia are part of the Australasian continent and have different germplasm biodiversity than the western islands. The area inside of the Wallacea Webber lines holds various endemic species. In the past, research by Hoogstral, Anastos, and their colleagues contributed significantly to our understanding of tick biodiversity in Indonesia, with more than 55 tick species reported to infest different animals in the region. Some of them have particular endemic hosts, such as Amblyomma robinsoni of Varanus komodoensis, Amblyomma babirussae of Babirussa babyrussa, Aponomma komodoense of Varanus komodoensis, and Amblyomma soembawensi of Varanus salvator [171][131]. The most frequently reported ticks affecting livestock and companion animals include R. sanguineus s.l., several clades of R. microplus (that may result in several species after adequate studies), Rhipicephalus pilans, and Haemaphysalis bispinosa [172,173,174][132][133][134]. Tick-borne diseases are reported mainly from highly populated areas like Java and Bali. Tick-borne pathogens in companion animals are Ehrlichia sp., Babesia sp., and Anaplasma sp. whereas livestock are infected by Babesia bigemina, B. bovis, Babesia naoakii, Theileria orientalis, Theileria sp., A. marginale, and Coxiella burnetti [174,175,176,177,178,179][134][135][136][137][138][139]. Tick-borne diseases pose a significant growing threat, mainly due to human activity and current climate trends. Recently, there has also been an upsurge in the number of tourists visiting remote islands in the region, leading to increased interest in these areas beyond Bali and Lombok. While the recent development progress in these regions benefits the nation’s growth, it poses a significant risk of exposure to ticks and tick-borne diseases from unknown areas that can be transmitted to new hosts. Although cases of ticks and tick-borne diseases are persistently reported, rural and urban societies pay less attention to them than mosquito-borne illnesses. 

2.12. Turkey

Turkey is situated at the intersection of Asia, Europe, and Africa. This unique position allows the inclusion of different climatic regions, habitat types, and animal diversity, all of which provide suitable conditions to harbor different tick species. Moreover, Turkey contains migration routes and breeding and wintering areas of many migratory birds which bring together the risk of introduction and establishment of different tick populations and associated pathogens [189,190][140][141]. More than 40 tick species have been reported in the country to date [191][142]. The most-recorded species were H. marginatum, Hyalomma excavatum, Hyalomma anatolicum, Hyalomma asiaticum, Hyalomma aegyptium, R. sanguineus s.l., Rhipicephalus turanicus, Rhipicephalus bursa, Haemaphysalis parva, and Dermacentor marginatus [192,193,194,195,196,197,198][143][144][145][146][147][148][149]. As a result of this tick species richness, a number of pathogens have been detected, including Ehrlichia canis, Theileria spp. (Theileria ovis, T. annulata), Anaplasma spp. (A. marginale, A. phagocitophylum, A. platys, A. ovis, A. centrale, A. bovis), Borrelia spp. (B. burgdorferi s.l., B. turcica), Babesia spp. (Babesia ovis, B. bovis, B. bigemina, Babebsia major, Babebsia crassa, Babebsia canis, and B. divergens), Rickettsia spp. (Rickettsia aeschlimannii, Rickettsia hoogstraali, Rickettsia barbariae), and Hepatozoon canis [199,200,201,202,203,204,205][150][151][152][153][154][155][156]. Furthermore, a considerable number of tick-borne viruses have been reported [206,207,208,209][157][158][159][160]. Lyme borreliosis and tick-borne encephalitis (TBE) are not prevalent in Turkey, although I. ricinus, the vector of these diseases, is widely distributed in the northern parts of the country [210,211,212,213][161][162][163][164]. CCHF constitutes a significant public health threat in Turkey since the first case was reported in 2002. Based on official records, 10,562 cases have been recorded from 2002–2017 and 501 of them resulted in death (https://hsgm.saglik.gov.tr/tr/zoonotikvektorel-kkka, accessed on 6 October 2023). Crimean-Congo hemorrhagic fever cases were mostly documented in rural areas in the central and northern regions of the country where agricultural and animal husbandry activities are common. Following the diagnosis of CCHF in Turkey, studies predominantly concentrated on the detection of CCHFV in ticks collected from these endemic regions [191,214,215,216,217,218][142][165][166][167][168][169]. 

2.13. Australia

Girt by sea, Australia has been in complete isolation for 40 million years. This separation has led to the evolution of 70 characterized species of argasid and ixodid ticks that have co-evolved with Australia’s unique mammalian (e.g., Ixodes ornithorynchi, platypus tick, and Amblyomma triguttatum, the ornate kangaroo tick), avian (e.g., Argas robertsi, Roberts’ bird tick), and reptilian (e.g., Amblyomma albolimbatum, stumpy-tailed lizard tick) fauna [225][170]. The most common biting ticks in Australia include I. holocyclus and A. triguttatum on the east and west coasts that parasitize people, respectively; Haemaphysalis longicornis and Rhipicephalus australis for cattle; and R. linnaei for dogs. It has been hypothesized that hard ticks evolved in the part of Gondwana that later became Australasia (~120 million years ago), evidenced by the basal lineage of Metastriata, Bothriocrotoninae, and Australian lineages of Ixodes, unique to Australia [226][171]. Despite Australia’s isolation, five tick species have been introduced into the Australian continent due to the movement of domestic animals following the arrival of Europeans in 1788 [227][172]. This introduction has led to the incursion of several tick-borne pathogens that affect companion and livestock animals in Australia, including A. platys, B. vogeli, Borrelia persica, T. orientalis complex, and more recently, H. canis and E. canis. Regarding human TBDs, only three are formally accepted: Queensland spotted fever, Flinders Island spotted fever, and Q fever [228][173]. The advancement of molecular techniques in recent years has led to the exponential discovery of several taxa of interest (TOIs; taxa closely related to known global tick-borne pathogens) identified within Australian ticks, wildlife, and domestic animals [229][174]. TOIs include Borrelia tachyglossi harbored within the echidna tick Bothriocroton concolor and several closely related unnamed Borrelia spp. in Bothriocroton undatum and within introduced and native rodents; a whole suite of Anaplasmatacae, Francisellaceae, Midichloriaceae, Coxiellaeceae, Bartonellaceae, Mycoplasmatcceae, and Rickettsiaceae species, including Neoehrlichia australis and Neoehrlichia arcana, Midichloria mitochondrii, Coxiella massiliensis, hemotropic mycoplasmas and novel species of Anaplasma, Ehrlichia, Rickettsia, Francisella; rhabdoviruses, chuviruses, coltivurses, flavivurses, and jingmenviruses; lastly, hemoprotozoa have also been recently discovered, including Thelieria spp., Babesia spp., Trypanosoma spp., and Hepatozoon spp. [229,230,231,232,233,234,235,236,237,238,239,240][174][175][176][177][178][179][180][181][182][183][184][185]. The genetic diversity of these TOIs mirrors the co-evolution of ticks and native wildlife. Furthermore, the uniqueness of these microbes answers why standard genus-specific and even species-specific assays from the northern hemisphere have failed in previous years to characterize tick-borne microbes in Australian ticks. 

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