Uma revisão sobre duas espécies distintas de Rhipicephalus: R. microplus e R. Australis

cr Ciência Rural Cienc. Rural 0103-8478 1678-4596 Universidade Federal de Santa Maria RESUMO: As espécies pertencentes ao gêneroRhipicephalus (Boophilus) são carrapatos monoxenos de distribuição sazonal em regiões tropicais e subtropicais. Por muito anos, duas espécies de carrapatos foram consideradas comoRhipicephalus microplus.Contudo, estudos recentes reclassificaram esse carrapato em duas espécies: R. micropluseR. australis. Em razão de diversas semelhanças entre R. micropluse R. australis,distinguir essas duas espécies torna-se uma tarefa árdua, o que explica as mudanças de classificação dessas espécies nas últimas décadas. O reaparecimento da espécieR. australissurge com novas pesquisas, resultado de uma melhor caracterização dessas prováveis espécies crípticas. Evidências baseadas em análises das características morfológicas, na ausência de co-especificidade, em marcadores de microssatélites, no DNA ribossomal mitocondrial 12S e 16S, assim como no genoma mitocondrial, suportam a re-classificação deR. micropluscomo duas espécies distintas. Nesse sentido, populações deR. microplusda Austrália, Camboja, Nova Caledônia, Bornéo, Filipinas, Nova Guiné, Indonésia e Taiti foram recentemente renomeadas comoR. australis. Além disso, um melhor entendimento sobre a especiação e localização dessas duas espécies pode trazer avanços importantes para melhorar as estratégias de controle desses carrapatos. INTRODUCTION: Records of ticks date back to approximately 2,000 years, when they were considered anus-less animals by Pliny, the Elder (A.D. 23-79) (HILLYARD, 1996). Later, reverend Dr. Thomas Moufet (A.D. 1553-1604) noted that Ricinus means "filled with food abundantly and yet there is no passage for any excrement" (BOWMAN & SAUER, 2004). To answer the question from "where the first ticks were evolved?" DOBSON & BARKER (1999) suggested that ticks evolved in the part of the supercontinent Gondwana that became Australia, in the early Devonian era, around 390 Mya ago. Contrarily, KLOMPEN et al. (2000) suggested that the hard ticks evolved 120 Mya later, after Australia became relatively isolated. Ticks are hematophagous acarines mainly distributed in two major families, Ixodidae(hard tick) and Argasidae (soft tick) (BLACK & PIESMAN, 1994), while only one tick species is present in the Nuttalliellidae family (GUGLIELMONE et al., 2010). Argasid ticks feed fast, but take in small volumes of blood in each meal, contrasting with Ixodid ticks, which have a longer-feeding process and ingest large amounts of blood. Ixodid ticks are unique among ectoparasites due to their long host attachment period (SAUER et al., 2000) and are divided into metastriate and prostriate ticks. Metastriate ticks include Rhipicephalus genera, characterized by relatively short mouthparts and secretion of copious amounts of cement or glue responsible for the firm attachment to its host. Prostriate ticks include the Ixodes genera, with longer and barbed mouthparts that are responsible for attachment to the host (FRANCISCHETTI et al., 2009). Studies demonstrated differences of specific Rhipicephalus(Boophilus) spp. life cycle in different ecological zones, where varying degrees of climatic conditions affect each phase of the tick life cycle (LEGG, 1930; LONDT & ARTHUR, 1975) and changes in these factors can difficult precise establishing of the life cycles of R. microplusand R. australis. In recent years, Boophilus spp. was reassigned to the genus Rhipicephalus(MURRELL & BARKER, 2003). Since, based on molecular and morphological studies, some Rhipicephalus species were found to be more closely associated to R. (Boophilus) species in comparison to Rhipicephalus species (MURRELL et al., 2000). Substantial morphological and molecular data have produced important evidence supporting the genus Rhipicephalus as paraphyletic to the genus Boophilus (MURRELL & BARKER, 2003; BARKER & MURRELL, 2004). To avoid misunderstanding and disbelief among researchers concerning name changes, Boophiluswas retained as a subgeneric epithet, and Boophilus microplus became Rhipicephalus(Boophilus) microplus(GUGLIELMONE et al., 2010) or Rhipicephalus microplus. The controversy about the classification of R. australis and R. microplusstarted in 1899, when the morphological description of Rhipicephalus australis moved this tick from the group of rhipicephalines species (FULLER, 1899). However, differences in morphological features, reproductive parameters, and genetic constitution between R. microplusspecimens collected in America and Africa and individuals collected in Australia provide sufficient evidences to reclassify R. australis and R. microplusas different species (LABRUNA et al., 2009; ESTRADA-PEÑA et al., 2012). Based on such evidence, we reviewed the data supporting the current taxonomic position of R. microplusand R. australis as distinct species. Current taxonomy Morphological differences R. microplus, earlier named Haemaphysalis micropla Canestrini, 1888, was not precisely distinguished from R. australis when this species was identified by Fuller (FULLER, 1899). However, SALMON & STILES (1901) classified R. australis as a distinct species, while NEUMANN (1901) regarded R. australis as a subspecies of R. microplus. Several studies initially supported the conspecificity between these two species (BEDFORD, 1932; ROBERTS, 1965). Overall differences in specimens collected in Australia, America, and Africa are allegedly strong enough to support R. australis as a distinct species (MINNING, 1934). Contrarily, UILENBERG (1962) concluded that these morphological variations did not validate the notion that R. australis is a distinct species; therefore the author synonymized it under the name B. microplus. Similarly, for LONDT & ARTHUR (1975) the morphological variation observed between Australian and South African R. microplusdo not afford to maintain these two tick populations as distinct species. Contrarily, recent studies suggest that both female and male adults of R. australis, besides this species' clearly smaller larva, differ from R. microplus by a combination of other morphological characters (ESTRADA-PEÑA et al., 2012). Despite the features closely shared by R. australis and R. microplus, a clear set of morphological differences persists. Larvae of R. australis described as B. microplus using specimens collected in Australia are smaller, with a narrow dorsal scutum than R. microplus(CLIFFORD et al., 1961; ESTRADA-PEÑA et al., 2012). Adult R. australis can be recognized by ventro-medial spurs in male palpal segments, and the copious, plumose, pale white setae on the female dorsum. Variability in other features like adanal and coxal shields among different populations often lead to incorrect classifications. R. australis possess abundant longer and pale dorsal setae, and the median alloscutal setae are arranged in clusters of 4 to 6 rows. However, this feature may be lost in engorged females. R. microplusdorsal setae are smaller and slim, and medial alloscutal setae are compose by clusters of 2 to 3 rows (WALKER et al., 2003). As compared to R. microplus, the medial scutal setae in R. australis female are longer. The setae behind the eyes are visible in R. australis female but unapparent R. microplusfemale. The spur in the ventral surface of R. australis male is absent in R. microplusmale. Finally, in R. microplusmales possess numerous setae on the lateral margins of the ventral surface of the capitulum, which are smaller in R. australis(WALKER et al., 2003; ESTRADA-PEÑA et al., 2012). Genetic differences Crosses mating trials Cross mating studies are a useful tool in the differentiation of tick species. For example, R. annulatus and R. microplusappear to be closely related within the subgenus Boophilus, as compared to other R. (Boophilus) species (BEATI & KEIRANS, 2001). R. microplusand R. annulatus cross-breeding has revealed that these two species lack conspecificity (GRAHAM et al., 1972; THOMPSON et al., 1981; DAVEY et al., 1983). A boundary in distributions of R. microplusand R. annulatus along the Texas-Mexico border suggests that these species reproduced without cross-breeding (LOHMEYER et al., 2011). In the same way, genetic differences among R. microplusstrains found at specific bio-geographical and ecological areas led taxonomists to consider boophilid ticks from America/Africa as different species against those from Australia. The first evidence supporting the lack of genetic conspecificity between African and Australian R. micropluswas provided by SPICKETT & MALAN (1978), regarding these species as 'diverging taxa' under speciation process with recent history as distinct populations. Conversely, GUGLIELMONE et al., (2003) concluded that R. microplusfrom South Africa and Australia should be considered as distinct species. Crossbreeding of Australian and Argentinean or Mozambican population of boophilid ticks has led to infertile animals, differently of Argentinean and Mozambican population crosses, that resulted in fertile offspring. These observations support the conclusion that tick population from Mozambique and Argentina represent a single species, while tick from Australia is possibly a distinct species (LABRUNA et al., 2009). Furthermore, mating experiments between R. microplusfrom South Africa and R. australis from Australia produced infertile offspring (SPICKETT & MALAN, 1978), which suggests reproductive isolation between these two species. Differences based on mitochondrial genome In addition to morphological observations, mitochondrial 12S and 16S ribosomal DNA (rDNA) has been considered a promising tool to determine phylogenetic relationships among various tick species (MANGOLD et al., 1998; BEATI & KEIRANS, 2001; LABRUNA et al., 2009; ESTRADA-PEÑA et al., 2012; McCOOKE et al., 2015). Indeed, phylogenetic analysis based on rDNA also supports the lack of conspecificity among ticks (ESTRADA-PEÑA et al., 2012). In this way, rDNA phylogenies using sequences from a number of Australian populations lend strength to suggestion that two species is classified under the name R. microplus. Besides the absence of consistent data on specimens and the few number of populations employed for the molecular and cross-breeding by LABRUNA et al. (2009), the authors believe that the available information is sufficient to support the taxonomic separation of R. australis and R. microplus. As discussed above, the phylogenetic relationships among R. (Boophilus) species were mostly carried out based on evolutionary relationships, relying on partial rDNA sequences (MURRELL et al., 2000, 2001, 2003; BEATI & KEIRANS, 2001; LABRUNA et al., 2009; ESTRADA-PEÑA et al., 2012). Molecular analysis (BEATI & KEIRANS, 2001) has confirmed the sister-species relationship between R. annulatus and R. microplus, as previously suggested based on morphology (FELDMAN-MUHSAM & SHECHTER, 1970). Phylogenetic evidence (MURRELL et al., 2001) about the genus Rhipicephalus also supports monophyly of the Boophilus clade; however, this evidence did not solve the relationships among R. (Boophilus) species. Moreover, the 'R. microplus' specimens used in that work were from Australia (R. australis), and did not include R. micropluss.s. Recent studies based on a wide geographic sampling range of R. microplusalso support monophyly of the subgenus Rhipicephalus(Boophilus), thus the reinstatement of R. australis (LABRUNA et al., 2009; ESTRADA-PEÑA et al., 2012). Intriguingly, R. microplusstrains collected in India and Nepal show high divergence from American and African R. microplusstrains (LABRUNA et al., 2009). However, the phylogenetic placement of Indian and Nepalese R. microplusstrains was not resolved, though Indian R. microplusclustered with R. annulatus in a 16S rRNA analysis (LABRUNA et al., 2009). Mitochondrial genomes were investigated in tick phylogenetic relationships, and have been proved useful in solving tick lineages (BURGER et al., 2012, 2013, 2014a,b; WILLIAMS-NEWKIRK et al., 2015; McCOOKE et al., 2015). To date, a few mitochondrial genomes of rhipicephaline ticks have been sequenced, and include R. sanguineus (BLACK & ROEHRDANZ, 1998), R. simus (Xu et al., 2014) and 13 gene fragments of R. australismitochondrial genome (CAMPBELL & BARKER, 1999). Whole and incomplete mitochondrial genomes of R. annulatus,R. australis,R. kohlsi, R. geigyi, and R. microplusstrains from Brazil, Texas (USA) Cambodia and China were recently sequenced (BURGER et al., 2013; McCOOKE et al., 2015). Mitochondrial, cox1 and 16S rRNA phylogenetic analyses showed a species complex of R. annulatus, R. australis, and two clades of R. microplus. Morphologically related China and India R. micropluspopulation were found closed to R. annulatus than other specimens of R. microplusfrom Asia, South America, and Africa. Furthermore, cox1 and 16S rRNA nucleotide sequences afforded to resolve the phylogenetic relationships within the R. micropluscomplex more successfully, as compared to 12S rRNA or the nuclear marker ITS2 (CAMPBELL & BARKER, 1999; LEMPEREUR et al., 2010; BURGER et al., 2013). Microsatellite markers The genetic differences between R. micropluspopulations from the TEQA (tick eradication quarantine area) USA, Mexico, and Puerto Rico were first investigated using isozymes over three decades ago (SATTLER et al., 1986). Analysis of fifteen selected polymorphic loci demonstrated heterozygosity to arthropods and high genetic similarity among various strains, suggesting an undifferentiated gene pool in R. micropluspopulations prevalent in North America. Later, microsatellite polymorphism was investigated in tick populations from Argentina, Australia, India, Mozambique, and New Caledonia. Tissues of homologous and heterologous crosses were analyzed, and allele numbers and sizes were determined for each locus. The clustering analysis of microsatellite alleles from R. microplusoffspring crosses revealed that Australian homologous and heterologous crosses had lower similarity, in comparison to homologous and heterologous crosses between Mozambique and Argentinian strains (LABRUNA et al., 2009). In New Caledonia, R. microplushas been found in close contact with sympatric hosts, on cattle and on a new host, the rusa deer (Cervus timorensis), divided into two differentiated genetic pools. In North America, the white-tailed deer (Odocoileus virginianus) and the red deer are also well-studied examples that illustrate this pattern of new sympatric hosts. This phenomenon may have occurred across the globe, in areas where this tick has established some degree of interaction with distinct host species in sympatry with livestock (KOFFI et al., 2006a; DE MEEUS et al., 2010; ARAYA-ANCHETTA et al., 2015). Using eight microsatellite loci, low genetic structure was found in specimens of R. australis in New Caledonia (KOFFI et al., 2006b). Analyses using 11 microsatellite markers revealed that the genetic differences between AustralianR. australis populations endemic in Queensland and New South Wale are not significant; however, the observed diversity among strains within each location proposes limitations to local gene flow (CUTULLÉ et al., 2009). A significant and small genetic structure was detected in ticks that infest rusa deer or domestic cattle (De MEEUS et al., 2010), which suggests that this sympatric adaptation resulted in host-specific populations. Recently, microsatellite markers were successfully used to detect genetic variation among R. microplusstrains in Texas (BUSCH et al., 2014). Undifferentiated tick collections were analyzed using 11 repeat loci, and it was not possible to identify any genetic divergence between R. microplussampled from white-tailed deer and specimens collected from cattle, even when the ticks were obtained from the same pastures (BUSCH et al., 2014). When parasitized by cattle ticks, these wild ungulates pose a major obstacle to eradication programs, since the difficulty to treat these animals with acaricides (POUND et al., 2010). Differences in Bm86 gene sequences Characterization employing an intron and DNA short tandem repeats (STRs) from Bm86 gene of Latin America and Australia R. micropluspopulations revealed significant differences within the Bm86 coding region between ticks from these regions (De La FUENTE et al., 2000). The intron sequence dissipated in the phylogenetic analysis, showing that individuals of the same strain do indeed vary. Moreover, polymorphism analysis at STR suggests differences within and between populations of R. microplus, which supports the molecular level of existence between R. micropluspopulations (De La FUENTE et al., 2000). Ecological niche Field data on regional distribution are necessary to clarify the actual distribution and the evolution of R. microplusandR. australis. It has been hypothesized that R. microplusoriginated in South and Southeast Asia (HOOGSTRAAL, 1985), later spreading to Madagascar and Southern Africa. Data required to shed more light on how and when R. microplusspread to Americas is scarce (LABRUNA et al., 2009), although it is reasonable that the tick was brought attached on livestock imported from India or Africa 4-5 centuries ago (BARRÉ & UILENBERG, 2010). Since R. microplusis specific to ungulates, the transportation of European cattle (Bos taurus) has spread it throughout the tropical and subtropical belt. As compared to tropical Bovidae, European bovines are almost incapable of eliciting efficient immune responses to R. microplusinfestations (FRISCH, 1999). Other factors that may have constrained the geographical distribution of R. microplusinclude climate variables and competition with other tick species (ESTRADA-PEÑA et al., 2006; CHEVILLON et al., 2013). Furthermore, it was suggested that R. australis was introduced in Australia and New Caledonia on livestock in 1829 (ANGUS, 1996), and 1942 (VERGES, 1944; DE MEEÛS et al., 2010), respectively. Today, R. australis is prevalent in Australia, Cambodia, Philippines, Indonesia, New Caledonia, Borneo, Malaysia, New Guinea and Tahiti (LABRUNA et al., 2009; ESTRADA-PENA et al., 2012; LOW et al., 2015). R. microplusis endemic between parallels 32°N and 32°S, where the major cattle breeding countries are, including newly reported ecological preferences in West Africa (MADDER et al., 2007; ESTRADA-PEÑA et al., 2006; LEGER et al., 2013). In addition, R. microplusand R. australis coexist in southeastern Asian countries (LOW et al., 2015). However, climate changes may further modify the potential geographical distribution of these parasites. Implications in control methods Economically, ticks are among the most potentially harmful parasites worldwide (PIESMAN & EISEN, 2008; GRISI et al., 2014). During infestation, ticks may transmit numerous pathogenic bacterial, viral, and protozoan organisms to the host (HAJDUSEK et al., 2013). The combination of the tick vectoring ability and the overall undermining of the host's health caused by blood uptake leads to high losses in livestock and pet industries (WILLADSEN, 2004). R. micropluswas introduced in the New World by tick-infested cattle brought by explorers and colonists (HOOGSTRAAL, 1985). Estimated losses caused by these ectoparasites during the first decade of the 19th century reached about US$63 billion (MOHLER, 1906). Therefore, due to the huge economic impact caused by R. microplus, the United States started a national campaign to eradicate Boophilus in 1906, and by 1943 this program was declared complete (GRAHAM & HOURRIGAN, 1977). Considering Brazil alone, which has the largest industrial cattle herd in the world; potential losses caused by R. microplusare estimated at US$3.24 billion yearly (GRISI et al., 2014). Tick control is a daunting challenge, because current control methods based on synthetic acaricides are becoming increasingly inefficient (RECK et al., 2014). Effective new control strategies demand a precisely parasitological epidemiology, because these strategies are affected by the presence ofR. microplusin a given geographical area. Vaccination experiments demonstrated different degrees of susceptibility for Bm86-based vaccines by R. micropluspopulations, suggesting the existence of genetically distinct strains of R. microplus(COBON et al., 1995; GARCIA-GARCIA et al., 2000). The Bm86 vaccine developed from an Australian 'R. microplus' strain showed very low efficacy when used in Latin America, as compared to Bm95, a Bm86 homologue from the Argentinian R. microplusstrain, which showed higher vaccinal efficacy in this region (GARCIA-GARCIA et al., 2000). Likewise, a Bm86 formulation from Cuban R. microplusshowed higher protection levels in America, when compared with Bm86 from Australian 'R. microplus' (De La FUENTE et al., 2007). Consequently, it is important to clearly describe vaccination data regarding the tick strain analyzed, due to the observed species-specificity of protection levels. Moreover, risk areas for tick-borne disease also need re-analysis, given the potential differences in vector capability by these two tick species. CONCLUSION: The triumph of taxonomy at species level is an open tool for continuous and exciting discoveries. Better understanding of R. microplusand R. australis taxonomic structure is pivotal for control measures against these ticks. Further studies on population genetic of Rhipicephalus(Boophilus) spp. will more thoroughly explain the interactions among these parasites, the pathogens they vector, and their hosts. In addition, such studies will provide more in-depth information about tick movements, disease dispersal, and designing anti-tick control programs. Genetic variance studies among Rhipicephalus(Boophilus) spp. will better elucidate the identification of cryptic species. Mixed infestation, when the parasites are in close contact with sympatric alternative hosts (like deer, for instance) is a major hurdle for eradication programs. However, more detailed information about genetic differentiation and genetic population structure could provide tools to improve control strategies. As demonstrated in recent years, the lack of effective protection induced by the Bm86 vaccine can be partly explained considering the presence of different R. microplusstrains. These failed vaccine experiments can be explained if R. microplusis considered not a single, but two distinct species. Although R. australis is very similar to R. microplus, both the larvae and adults of each species have a clear set of species-specific characters. In conclusion, reported results confirm that two distinct species of ticks were named R. microplus. ACKNOWLEDGMENT This work was supported by grants from Higher Education Commission, Pakistan Science Foundation (Pakistan), Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq), and Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES). We would like to thanks Dr. Carlos Termignoni for critical reading of this review. REFERENCES: ALI, A. et al. Probing the functional role of tick metalloproteases. Physiological Entomology, v.40, p.177, 188, 2015. Available from: <Available from: http://onlinelibrary.wiley.com/doi/10.1111/phen.12104/abstract >. Accessed: Sept. 27, 2015. doi: 10.1111/phen.12104. ALI A. Probing the functional role of tick metalloproteases Physiological Entomology 40 177 188 177, 188 2015 Available from: http://onlinelibrary.wiley.com/doi/10.1111/phen.12104/abstract Sept. 27, 2015 10.1111/phen.12104. ANGUS, B.M. The history of the cattle tick Boophilus microplus in Australia and achievements in its control. International Journal for Parasitology, v.26, p.1341-1355, 1996. Available from: <Available from: http://www.sciencedirect.com/science/article/pii/S0020751996001129 >. Accessed: Jun. 05, 2015. doi: 10.1016/s0020-7519(96)00112-9. ANGUS B.M. The history of the cattle tick Boophilus microplus in Australia and achievements in its control. International Journal for Parasitology 26 1341 1355 1996 Available from: http://www.sciencedirect.com/science/article/pii/S0020751996001129 Jun. 05, 2015 10.1016/s0020-7519(96)00112-9. ARAYA-ANCHETTA, A. et al. Thirty years of tick population genetics: A comprehensive review. Infection Genetics and Evolution, v.29, p.164-179, 2015. Available from: <Available from: http://www. sciencedirect.com/science/article/pii/S1567134814004110 >. Accessed: Oct. 01, 2015. doi: 10.1016/j.meegid.2014.11.008. ARAYA-ANCHETTA A. Thirty years of tick population genetics: A comprehensive review. Infection Genetics and Evolution 29 164 179 2015 Available from: http://www. sciencedirect.com/science/article/pii/S1567134814004110 Oct. 01, 2015 10.1016/j.meegid.2014.11.008 BARKER, S.C.; MURRELL, A. Systematics and evolution of ticks with a list of valid genus and species names. Parasitology, v.129, p.S15-S36, 2004. Available from: <Available from: http://www.ncbi.nlm.nih.gov/pubmed/15938503 >. Accessed: Feb. 07, 2015. doi: 10.1017/S0031182004005207. BARKER S.C. MURRELL A. Systematics and evolution of ticks with a list of valid genus and species names. Parasitology 129 S15 S36 2004 Available from: http://www.ncbi.nlm.nih.gov/pubmed/15938503 Feb. 07, 2015 10.1017/S0031182004005207. BARRÉ, N.; UILENBERG, G. Spread of parasites transported with their hosts: case study of two species of cattle tick. Revue scientifique et technique (International Office of Epizootics), v.29, p.149-160, 2010. BARRÉ N. UILENBERG G. Spread of parasites transported with their hosts: case study of two species of cattle tick. Revue scientifique et technique (International Office of Epizootics 29 149 160 2010 BEATI, L.; KEIRANS, J.E. Analysis of the systematic relationships among ticks of the genera Rhipicephalus and Boophilus (Acari: Ixodidae) based on mitochondrial 12S ribosomal DNA gene sequences and morphological characters. Journal of, Parasitology v.87, p.32-48, 2001. Available from: <Available from: http://www.ncbi.nlm.nih.gov/pubmed/11227901 >. Accessed: Mar. 12, 2015. doi: 10.1645/0022-3395(2001)087[0032:AOTSRA]2.0.CO;2. BEATI L. KEIRANS J.E. Analysis of the systematic relationships among ticks of the genera Rhipicephalus and Boophilus (Acari: Ixodidae) based on mitochondrial 12S ribosomal DNA gene sequences and morphological characters Journal of, Parasitology 87 32 48 2001 Available from: http://www.ncbi.nlm.nih.gov/pubmed/11227901 Mar. 12, 2015 10.1645/0022-3395(2001)087[0032:AOTSRA]2.0.CO;2. BEDFORD, G.A.H. Asynoptic check-list and host-list of the ectoparasites found on South African Mammalia, Aves, and Reptilia. Journal of Veterinary Science and Animal Industry, v.7, p.69-110, 1932. BEDFORD G.A.H. Asynoptic check-list and host-list of the ectoparasites found on South African Mammalia, Aves, and Reptilia Journal of Veterinary Science and Animal Industry 7 69 110 1932 BLACK, W.C.; PIESMAN, J. Phylogeny of hard- and soft-tick taxa (Acari: Ixodidae) based on mitochondrial 16S rDNA sequences. Proceedings of the National Academy of Sciences USA, v.91, p.10034-10038, 1994. BLACK W.C. PIESMAN J. Phylogeny of hard- and soft-tick taxa (Acari: Ixodidae) based on mitochondrial 16S rDNA sequences. Proceedings of the National Academy of Sciences USA 91 10034 10038 1994 BLACK, W.C.; ROEHRDANZ, R.L. Mitochondrial gene order is not conserved in arthropods: prostriate and metastriate tick mitochondrial genomes. Molecular Biology and Evolution, v.15, p.1772-1785, 1998. BLACK W.C. ROEHRDANZ R.L. Mitochondrial gene order is not conserved in arthropods: prostriate and metastriate tick mitochondrial genomes. Molecular Biology and Evolution 15 1772 1785 1998 BOWMAN, A.S.; SAUER J.R. Tick salivary glands: function, physiology and future., Parasitology v.129, p.S67-S81, 2004. Available from: <Available from: http://www.ncbi.nlm.nih.gov/pubmed/15938505 >. Accessed: Apr. 2, 2015. doi: 10.1017/S0031182004006468. BOWMAN A.S. SAUER J.R. Tick salivary glands: function, physiology and future. Parasitology 129 S67 S81 2004 Available from: http://www.ncbi.nlm.nih.gov/pubmed/15938505 Apr. 2, 2015 10.1017/S0031182004006468. BURGER, T.D. et al. Molecular phylogeny of soft ticks (Ixodida: Argasidae) inferred from mitochondrial genome and nuclear rRNA gene sequences. Ticks and Tick-Borne Diseases, v.5, p.195-207, 2014b. Available from: <Available from: http://www.sciencedirect.com/science/article/pii/S1877959X13001234 >. Accessed: Jul. 14, 2015. doi: 10.1016/j.ttbdis.2013.10.009. BURGER T.D. Molecular phylogeny of soft ticks (Ixodida: Argasidae) inferred from mitochondrial genome and nuclear rRNA gene sequences Ticks and Tick-Borne Diseases 5 195 207 2014b Available from: http://www.sciencedirect.com/science/article/pii/S1877959X13001234 Jul. 14, 2015 10.1016/j.ttbdis.2013.10.009. BURGER, T.D. et al. Phylogenetic analysis of mitochondrial genome sequences indicates that the cattle tick, Rhipicephalus (Boophilus) microplus, contains a cryptic species. Molecular Phylogenetics and Evolution, v.76, p.241-253, 2014a. Available from: <Available from: http://www.sciencedirect.com/science/article/pii/S1055790314001171 >. Accessed: May 14, 2015. doi: 10.1016/j.ympev.2014.03.017. BURGER T.D. Phylogenetic analysis of mitochondrial genome sequences indicates that the cattle tick, Rhipicephalus (Boophilus) microplus, contains a cryptic species Molecular Phylogenetics and Evolution 76 241 253 2014a Available from: http://www.sciencedirect.com/science/article/pii/S1055790314001171 May 14, 2015 10.1016/j.ympev.2014.03.017. BURGER, T.D. et al. Phylogenetic analysis of the mitochondrial genomes and nuclear rRNA genes of ticks reveals a deep phylogenetic structure within the genus Haemaphysalis, and further elucidates the polyphyly of the genus Amblyomma with respect to Amblyomma sphenodonti and Amblyomma elaphense., Ticks and Tick-Borne Diseases v.4, p.265-274, 2013. Available from: <Available from: http://www.sciencedirect.com/science/article/pii/S1877959X13000344 >. Accessed: Apr. 06, 2015. doi: 10.1016/j.ttbdis.2013.02.002. BURGER T.D. Phylogenetic analysis of the mitochondrial genomes and nuclear rRNA genes of ticks reveals a deep phylogenetic structure within the genus Haemaphysalis, and further elucidates the polyphyly of the genus Amblyomma with respect to Amblyomma sphenodonti and Amblyomma elaphense. Ticks and Tick-Borne Diseases 4 265 274 2013 Available from: http://www.sciencedirect.com/science/article/pii/S1877959X13000344 Apr. 06, 2015 10.1016/j.ttbdis.2013.02.002 BURGER, T.D. et al. Phylogenetic analysis of ticks (Acari: Ixodidae) using mitochondrial genomes and nuclear rRNA genes indicates that the genus Amblyomma is polyphyletic. Molecular Phylogenetic and Evolution, v.64, p.45-55, 2012. Available from: <Available from: http://www.sciencedirect.com/science/article/pii/S1055790312001091 >. Accessed: Sept. 04, 2015. doi: 10.1016/j.ympev.2012.03.004. BURGER T.D. Phylogenetic analysis of ticks (Acari: Ixodidae) using mitochondrial genomes and nuclear rRNA genes indicates that the genus Amblyomma is polyphyletic Molecular Phylogenetic and Evolution 64 45 55 2012 Available from: http://www.sciencedirect.com/science/article/pii/S1055790312001091 Sept. 04, 2015 10.1016/j.ympev.2012.03.004 BUSCH, J.D. et al. Widespread movement of invasive cattle fever ticks (Rhipicephalus microplus) in southern Texas and shared local infestations on cattle and deer. Parasites & Vectors, v.7, p.188, 2014. Available from: <Available from: http://parasitesandvectors.biomedcentral.com/ >. Accessed: May 17, 2015. doi: 10.1186/1756-3305-7-188. BUSCH J.D. Widespread movement of invasive cattle fever ticks (Rhipicephalus microplus) in southern Texas and shared local infestations on cattle and deer Parasites & Vectors 7 188 188 2014 Available from: http://parasitesandvectors.biomedcentral.com/ May 17, 2015 10.1186/1756-3305-7-188. CAMPBELL, N.J.H.; BARKER, S.C. The novel mitochondrial gene arrangement of the cattle tick, Boophilus microplus: five fold tandem repetition of a coding region., Molecular Biology and Evolution v.16, p.732-740, 1999. CAMPBELL N.J.H. BARKER S.C. The novel mitochondrial gene arrangement of the cattle tick, Boophilus microplus: five fold tandem repetition of a coding region Molecular Biology and Evolution 16 732 740 1999 CHEVILLON, C. et al. Understanding the genetic, demographical and/or ecological processes at play in invasions: lessons from the southern cattle tick Rhipicephalus microplus (Acari: Ixodidae). Experimental and Applied Acarology, v.59, p.203-218. 2013. Available from: <Available from: http://link.springer.com/article/10.1007% 2Fs10493-012-9602-5 >. Accessed: Jun. 14, 2015. doi: 10.1007/s1 0493-012-9602-5. CHEVILLON C. Understanding the genetic, demographical and/or ecological processes at play in invasions: lessons from the southern cattle tick Rhipicephalus microplus (Acari: Ixodidae) Experimental and Applied Acarology 59 203 218 2013 Available from: http://link.springer.com/article/10.1007% 2Fs10493-012-9602-5 Jun. 14, 2015 10.1007/s1 0493-012-9602-5 CLIFFORD, C.M. et al. The larval Ixodid ticks of the eastern United States (Acarina: Ixodidae). Miscellaneous publications of the Entomological Society of America, v.2, p.213-237, 1961. CLIFFORD C.M. The larval Ixodid ticks of the eastern United States (Acarina: Ixodidae Miscellaneous publications of the Entomological Society of America 2 213 237 1961 COBON, G. et al. Vaccination against Boophilus microplus: the Australian field experience. In: FUENTE, J. de la (Ed.). Recombinant vaccines for the control of cattle tick. Havana: Elfos Scintiae, 1995. p.163-176. COBON G. Vaccination against Boophilus microplus: the Australian field experience. FUENTE J. de la Recombinant vaccines for the control of cattle tick Havana Elfos Scintiae 1995 163 176 CUMMING, G.S.; VUUREN D.V.P. Will climate change affect ectoparasite species ranges. Global Ecology and Biogeography, v.15, p.486-497, 2006. Available from: <Available from: http://onlinelibrary.wiley.com/doi/10.1111/j.1466-822X.2006.00241.x/abstract >. Accessed: May 15, 2015. doi: 10.1111/j.1466-822 X.2006.00241.x. CUMMING G.S. VUUREN D.V.P. Will climate change affect ectoparasite species ranges. Global Ecology and Biogeography 15 486 497 2006 Available from: http://onlinelibrary.wiley.com/doi/10.1111/j.1466-822X.2006.00241.x/abstract May 15, 2015 10.1111/j.1466-822 X.2006.00241.x CUTULLÉ, C. et al. Population structure of Australian isolates of the cattle tick Rhipicephalus (Boophilus) microplus. Veterinary, Parasitology v.161, p.283-291, 2009. Available from: <Available from: http://www.sciencedirect.com/science/article/pii/S0304401709000326 >. Accessed: Sept. 25, 2015. doi: 10.1016/j.vetpar.2009.01.005. CUTULLÉ C. Population structure of Australian isolates of the cattle tick Rhipicephalus (Boophilus) microplus. Veterinary, Parasitology 161 283 291 2009 Available from: http://www.sciencedirect.com/science/article/pii/S0304401709000326 Sept. 25, 2015 10.1016/j.vetpar.2009.01.005. DAVEY, R.B. et al. Longevity and mating behavior in males and parthenogenesis in females in hybridized Boophilus ticks (Acari: Ixodidae). Journal of Medical Entomology, v.20, p.614-617, 1983. DAVEY R.B. Longevity and mating behavior in males and parthenogenesis in females in hybridized Boophilus ticks (Acari: Ixodidae). Journal of Medical Entomology 20 614 617 1983 DE LA FUENTE , J. et al. A ten-year review of commercial vaccine performance for control of tick infestations on cattle. Animal Health Research Reviews, v.8, p.23-38, 2007. Available from: <Available from: http://www.ncbi.nlm.nih.gov/pubmed/17692140 >. Accessed: May 02, 2015. doi: 10.10 17/S1466252307001193. DE LA FUENTE J. A ten-year review of commercial vaccine performance for control of tick infestations on cattle. Animal Health Research Reviews 8 23 38 2007 Available from: http://www.ncbi.nlm.nih.gov/pubmed/17692140 May 02, 2015 10.10 17/S1466252307001193 DE LA FUENTE , J. et al. Molecular analysis of Boophilus spp. (Acari: Ixodidae) tick strains. Veterinary, Parasitology v.92, p.209-222, 2000. Available from: <Available from: http://www.ncbi.nlm.nih.gov/pubmed/10962158 >. Accessed: May 12, 2015. doi: 10.1016/S0304-4017(00)00308-3. DE LA FUENTE J. Molecular analysis of Boophilus spp. (Acari: Ixodidae) tick strains. Veterinary, Parasitology 92 209 222 2000 Available from: http://www.ncbi.nlm.nih.gov/pubmed/10962158 May 12, 2015 10.1016/S0304-4017(00)00308-3. DE MEEÛS, T. et al. Swift sympatric adaptation of a species of cattle tick to a new deer host in New Caledonia. Infection, Genetics and Evolution, v.10, p.976-983, 2010. Available from: <Available from: http://www.sciencedirect.com/science/article/pii/S1567134810001693 >. Accessed: Apr. 16, 2015. doi: 10.1016/j.meegid.2010.06.005. DE MEEÛS T. Swift sympatric adaptation of a species of cattle tick to a new deer host in New Caledonia. Infection, Genetics and Evolution 10 976 983 2010 Available from: http://www.sciencedirect.com/science/article/pii/S1567134810001693 Apr. 16, 2015 10.1016/j.meegid.2010.06.005. DOBSON, S.J.; BARKER, S.C. Phylogeny of the hard ticks (Ixodidae) inferred from 18S rRNA indicates that the genus Aponomma is paraphyletic., Molecular Phylogenetics and Evolution v.11, p.288-295, 1999. Available from: <Available from: http://www.ncbi.nlm.nih.gov/pubmed/10191073 >. Accessed: Jun. 28, 2015. doi: 10.1006/mpev.1998.0565. DOBSON S.J. BARKER S.C. Phylogeny of the hard ticks (Ixodidae) inferred from 18S rRNA indicates that the genus Aponomma is paraphyletic., Molecular Phylogenetics and Evolution 11 288 295 1999 Available from: http://www.ncbi.nlm.nih.gov/pubmed/10191073 Jun. 28, 2015 10.1006/mpev.1998.0565. ESTRADA-PEÑA, A. et al. Reinstatement of Rhipicephalus (Boophilus) australis (Acari: Ixodidae) with redescription of the adult and larval stages., Journal of Medical Entomology v.49, n.4, p.794-802, 2012. Available from: <Available from: http://www.ncbi.nlm.nih.gov/pubmed/22897039 >. Accessed: Jan. 13, 2015. doi: 10.1603/ME11223. ESTRADA-PEÑA A. Reinstatement of Rhipicephalus (Boophilus) australis (Acari: Ixodidae) with redescription of the adult and larval stages Journal of Medical Entomology 49 4 794 802 2012 Available from: http://www.ncbi.nlm.nih.gov/pubmed/22897039 Jan. 13, 2015 10.1603/ME11223. ESTRADA-PEÑA, A. et al. The known distribution and ecological preferences of the tick subgenus Boophilus (Acari: Ixodidae) in Africa and Latin America., Experimental and Applied Acarology v.38, p.219-235, 2006. ESTRADA-PEÑA A. The known distribution and ecological preferences of the tick subgenus Boophilus (Acari: Ixodidae) in Africa and Latin America Experimental and Applied Acarology 38 219 235 2006 FELDMAN-MUHSAM, B.; SHECHTER, R. Some notes on the genus Boophilus (Ixodidae) with special reference to species found in Israel., Journal of Medical Entomology v.7, p.677-686, 1970. Available from: <Available from: http://www.ncbi.nlm.nih.gov/pubmed/5501234 >. Accessed: Sept. 13, 2014. doi: 10.1093/jmedent/7.6.677. FELDMAN-MUHSAM B. SHECHTER R. Some notes on the genus Boophilus (Ixodidae) with special reference to species found in Israel Journal of Medical Entomology 7 677 686 1970 Available from: http://www.ncbi.nlm.nih.gov/pubmed/5501234 Sept. 13, 2014 10.1093/jmedent/7.6.677. FRANCISCHETTI, I.M. et al. The role of saliva in tick feeding. Frontier Bioscience (Landmark Ed) , v.14, p.2051-2088, 2009. FRANCISCHETTI I.M. The role of saliva in tick feeding. Frontier Bioscience (Landmark Ed) 14 2051 2088 2009 FRISCH, J. Towards a permanent solution for controlling cattle tick., International Journal for Parasitology v.29, p.57-71, 1999. Available from: <Available from: http://www.sciencedirect.com/science/article/pii/S0020751998001775 >. Accessed: Sept. 12, 2014. doi: 10.1016/S0020-7519(98)00177-5. FRISCH J. Towards a permanent solution for controlling cattle tick International Journal for Parasitology 29 57 71 1999 Available from: http://www.sciencedirect.com/science/article/pii/S0020751998001775 Sept. 12, 2014 10.1016/S0020-7519(98)00177-5. FULLER, C. Notes on the Queensland cattle tick and its relationship to the red water tick and the blue tick of Cape Colony (South Africa). Queensland Agricultural Journal, v.4, p.389-394, 1899. FULLER C. Notes on the Queensland cattle tick and its relationship to the red water tick and the blue tick of Cape Colony (South Africa) Queensland Agricultural Journal 4 389 394 1899 GARCIA-GARCIA, J.C. et al. Sequence variations in the Boophilus microplus Bm86 locus and implications for immunoprotection in cattle vaccinated with this antigen., Experimental and Applied Acarology v.23, p.883-895, 1999. GARCIA-GARCIA J.C. Sequence variations in the Boophilus microplus Bm86 locus and implications for immunoprotection in cattle vaccinated with this antigen Experimental and Applied Acarology 23 883 895 1999 GRAHAM, O.H. et al. Cross-mating experiments with Boophilus annulatus and B. microplus (Acarina: Ixodidae)., Journal of Medical Entomology v.9, p.531-537, 1972. Available from: <Available from: http://jme.oxfordjournals.org/content/9/6/531.long >. Accessed: Jan. 15, 2014. doi: 10.1093/jmedent/9.6.531. GRAHAM O.H. Cross-mating experiments with Boophilus annulatus and B. microplus (Acarina: Ixodidae) Journal of Medical Entomology 9 531 537 1972 Available from: http://jme.oxfordjournals.org/content/9/6/531.long Jan. 15, 2014 10.1093/jmedent/9.6.531. GRAHAM, O.H.; HOURRIGAN, J.L. Eradication programs for the arthropod parasites of livestock., Journal of Medical Entomology v.13, p.629-658, 1977. Available from: <Available from: http://www.ncbi.nlm.nih.gov /pubmed/328882 >. Accessed: May 12, 2014. doi: 10.1093/ jmedent/13.6.629. GRAHAM O.H. HOURRIGAN J.L. Eradication programs for the arthropod parasites of livestock Journal of Medical Entomology 13 629 658 1977 Available from: http://www.ncbi.nlm.nih.gov /pubmed/328882 May 12, 2014 10.1093/ jmedent/13.6.629 GRISI, L. et al. Reassessment of the potential economic impact of cattle parasites in Brazil. Revista Brasileira de Parasitologia Veterinaria, v.23, p.150-156, 2014. Available from: <Available from: http://www.ncbi.nlm.nih.gov/pubmed/25054492 >. Accessed: May 25, 2015. doi: 10.1590/S1984-29612014042. GRISI L. Reassessment of the potential economic impact of cattle parasites in Brazil. Revista Brasileira de Parasitologia Veterinaria 23 150 156 2014 Available from: http://www.ncbi.nlm.nih.gov/pubmed/25054492 May 25, 2015 10.1590/S1984-29612014042. GUGLIELMONE, A. et al. Ticks (Acari: Ixodidae) of the Neotropical Zoogeographic Region. The Netherlands: Atalanta Houten, 2003. (Special Publication, International Consortium on) Ticks and Tick-Borne Diseases. GUGLIELMONE A. Ticks (Acari: Ixodidae) of the Neotropical Zoogeographic Region The Netherlands Atalanta Houten 2003 Ticks and Tick-Borne Diseases GUGLIELMONE, A.A. et al. The Argasidae, Ixodidae and Nuttalliellidae (Acari: Ixodidae) of the world: a list of valid species names. Zootaxa, v.2528, p.1-28, 2010. Available from: <Available from: http://www.mapress.com/zootaxa/2010/f/z02528p028f.pdf >. Accessed: Jan. 23, 2014. GUGLIELMONE A.A. The Argasidae, Ixodidae and Nuttalliellidae (Acari: Ixodidae) of the world: a list of valid species names. Zootaxa 2528 1 28 2010 Available from: http://www.mapress.com/zootaxa/2010/f/z02528p028f.pdf Jan. 23, 2014 HAJDUSEK, O. et al. Interaction of the tick immune system with transmitted pathogens. Frontiers in Cellular and Infection Microbiology, v.3, p.26, 2013. Available from: <Available from: http://www.ncbi.nlm.nih.gov/pubmed/23875177 >. Accessed: Mar. 23, 2014. doi: 10.3389/fcimb.2013.00026. HAJDUSEK O. Interaction of the tick immune system with transmitted pathogens. Frontiers in Cellular and Infection Microbiology 3 26 26 2013 Available from: http://www.ncbi.nlm.nih.gov/pubmed/23875177 Mar. 23, 2014 10.3389/fcimb.2013.00026. HILLYARD, P.D. Ticks of North-West Europe: keys and notes for identification of the species. Shrewsbury: Linnean Society of London and the Estuarine and Coastal Sciences Association by Field Studies Council, 1996. 178p. HILLYARD P.D. Ticks of North-West Europe: keys and notes for identification of the species Shrewsbury Linnean Society of London and the Estuarine and Coastal Sciences Association by Field Studies Council 1996 178 HOOGSTRAAL, H. Argasid and nuttalliellid ticks as parasites and vectors. Advances in, Parasitology v.24, p.135-238, 1985. Available from: <Available from: http://www.ncbi.nlm.nih.gov/pubmed/3904345 >. Accessed: May 23, 2014. doi: 10.1016/S0065-308X(08)60563-1. HOOGSTRAAL H. Argasid and nuttalliellid ticks as parasites and vectors. Advances in, Parasitology 24 135 238 1985 Available from: http://www.ncbi.nlm.nih.gov/pubmed/3904345 May 23, 2014 10.1016/S0065-308X(08)60563-1. KLOMPEN, J.S.H. et al. Systematics and biogeography of hard ticks, a total evidence approach. Cladistics, v.16, p.79-102, 2000. Available from: <Available from: http://www.sciencedirect.com/science/article/pii/S0748300799901263 >. Accessed: Nov. 19, 2014. doi: 10.1006/clad.1999.0126. KLOMPEN J.S.H. Systematics and biogeography of hard ticks, a total evidence approach Cladistics 16 79 102 2000 Available from: http://www.sciencedirect.com/science/article/pii/S0748300799901263 Nov. 19, 2014 10.1006/clad.1999.0126. KOFFI, B.B. et al. Characterization of polymorphic microsatellite loci within a young Boophilus microplus metapopulation. Molecular Ecology, v.6 p.502-504, Notes 2006b. Available from: <Available from: http://agritrop.cirad.fr/533705/ >. Accessed: May 12, 2015. doi: 10.1111/j.1471-8286.2006.01295.x. KOFFI B.B. Characterization of polymorphic microsatellite loci within a young Boophilus microplus metapopulation. Molecular Ecology 6 502 504 2006b Available from: http://agritrop.cirad.fr/533705/ May 12, 2015 10.1111/j.1471-8286.2006.01295.x. KOFFI, B.B. et al. Founder effects, inbreeding and effective sizes in the Southern cattle tick: the effect of transmission dynamics and implications for pest management., Molecular Ecology v.15, p.4603-4611, 2006a. Available from: <Available from: http://onlinelibrary.wiley.com/doi/10.1111/j.1365-294X.2006.03098.x/abstract >. Accessed: Jan. 16, 2015. doi: 10.1111/j.1365-294X.2006.03098.x. KOFFI B.B. Founder effects, inbreeding and effective sizes in the Southern cattle tick: the effect of transmission dynamics and implications for pest management. Molecular Ecology 15 4603 4611 2006a Available from: http://onlinelibrary.wiley.com/doi/10.1111/j.1365-294X.2006.03098.x/abstract Jan. 16, 2015 10.1111/j.1365-294X.2006.03098.x. LABRUNA, M.B. et al. Allopatric speciation in ticks: genetic and reproductive divergence between geographic strains of Rhipicephalus (Boophilus) microplus. BMC Evolutionary Biology, v.9, p.46-58, 2009. Available from: <Available from: http://www.biomedcentral.com/1471-2148/9/46 >. Accessed: Jan. 12, 2014. LABRUNA M.B. Allopatric speciation in ticks: genetic and reproductive divergence between geographic strains of Rhipicephalus (Boophilus) microplus. BMC Evolutionary Biology 9 46 58 2009 Available from: http://www.biomedcentral.com/1471-2148/9/46 Jan. 12, 2014 LEGER, E. et al. Changing distributions of ticks. causes and consequences., Experimental and Applied Acarology v.59, p.219-244, 2013. Available from: <Available from: http://link.springer.com/article/10.1007%2Fs10493-012-9615-0 >. Accessed: Dec. 23, 2014. doi: 10.1007/s10493-012-9615-0. LEGER E. Changing distributions of ticks. causes and consequences. Experimental and Applied Acarology 59 219 244 2013 Available from: http://link.springer.com/article/10.1007%2Fs10493-012-9615-0 Dec. 23, 2014 10.1007/s10493-012-9615-0. LEGG, J. Some observations on the life history of the cattle tick (Boophilus australis). Proceeding of Royal Society Queensland, v.41, p.121-32, 1930. LEGG J. Some observations on the life history of the cattle tick (Boophilus australis) Proceeding of Royal Society Queensland 41 121 132 1930 LEMPEREUR, L. et al. Development and validation of a PCR-RFLP test to identify African Rhipicephalus (Boophilus) ticks. Acta Tropica, v.114, p.55-58, 2010. Available from: <Available from: http://www.sciencedirect.com/science/article/pii/S0001706X10000070 >. Accessed: Sept. 13, 2014. doi: 10.1016/j.actatropica.2010.01.004. LEMPEREUR L. Development and validation of a PCR-RFLP test to identify African Rhipicephalus (Boophilus) ticks. Acta Tropica 114 55 58 2010 Available from: http://www.sciencedirect.com/science/article/pii/S0001706X10000070 Sept. 13, 2014 10.1016/j.actatropica.2010.01.004. LOHMEYER, K.H. et al. Distribution of Rhipicephalus (Boophilus) microplus and Rhipicephalus (Boophilus) annulatus (Acari: Ixodidae) infestations detected in the United States along the Texas/Mexico Border., Journal of Medical Entomology v.48, p.770-774, 2011. Available from: <Available from: http://jme.oxfordjournals.org/content/48/4/770.long >. Accessed: Oct 16, 2015. doi: 10.1603/ME10209. LOHMEYER K.H. Distribution of Rhipicephalus (Boophilus) microplus and Rhipicephalus (Boophilus) annulatus (Acari: Ixodidae) infestations detected in the United States along the Texas/Mexico Border. Journal of Medical Entomology 48 770 774 2011 Available from: http://jme.oxfordjournals.org/content/48/4/770.long Oct 16, 2015 10.1603/ME10209. LONDT, J.G.H.; ARTHUR, D.R. The structure and parasitic life cycle of Boophilus microplus (Canestrini, 1888) in South Africa (Acarina: Ixodidae). Journal of the Entomological Society of South Africa, v. 38 p.321-340, 1975. LONDT J.G.H. ARTHUR D.R. The structure and parasitic life cycle of Boophilus microplus (Canestrini, 1888) in South Africa (Acarina: Ixodidae) Journal of the Entomological Society of South Africa 38 321 340 1975 LOW, V. et al. Molecular characterization of the tick Rhipicephalus microplus in Malaysia: new insights into the cryptic diversity and distinct genetic assemblages throughout the world., Parasites & Vectors v.8, p.341, 2015. Available from: <Available from: http://www.ncbi.nlm.nih.gov/pmc/articles/PMC4482097/ >. Accessed: May 16, 2015. doi: 10.1186/s13071-015-0956-5. LOW V. Molecular characterization of the tick Rhipicephalus microplus in Malaysia: new insights into the cryptic diversity and distinct genetic assemblages throughout the world. Parasites & Vectors 8 341 341 2015 Available from: http://www.ncbi.nlm.nih.gov/pmc/articles/PMC4482097/ May 16, 2015 10.1186/s13071-015-0956-5. MADDER, M. et al. Rhipicephalus (Boophilus) microplus ticks found in West Africa., Experimental and Applied Acarology v.43, p.233-234, 2007. Available from: <Available from: http://link.springer.com/article /10.1007%2Fs10493-007-9110-1 >. Accessed: May 27, 2014. doi: 10.1007/s10493-007-9110-1. MADDER M. Rhipicephalus (Boophilus) microplus ticks found in West Africa. Experimental and Applied Acarology 43 233 234 2007 Available from: http://link.springer.com/article /10.1007%2Fs10493-007-9110-1 May 27, 2014 10.1007/s10493-007-9110-1. MANGOLD, J.J. et al. Mitochondrial 16S rDNA sequences and phylogenetic relationships of species of Rhipicephalus and other tick genera among Metastriata (Acari: Ixodidae). ParasitologyResearch, v.84, p.478-484, 1998. Available from: <Available from: http://www.ncbi.nlm.nih.gov/pubmed/9660138 >. Accessed: Jan. 12, 2014. doi: 10.1007/s004360050433. MANGOLD J.J. Mitochondrial 16S rDNA sequences and phylogenetic relationships of species of Rhipicephalus and other tick genera among Metastriata (Acari: Ixodidae) Parasitology 84 478 484 1998 Available from: http://www.ncbi.nlm.nih.gov/pubmed/9660138 Jan. 12, 2014 10.1007/s004360050433. McCOOKE, J.K. et al. The mitochondrial genome of a Texas outbreak strain of the cattle tick, Rhipicephalus (Boophilus) microplus, derived from whole genome sequencing Pacific Biosciences and Illumina reads. Gene, v.571, p.135-141, 2015. Available from: <Available from: http://www.ncbi.nlm.nih.gov/pubmed/26117172 >. Accessed: Oct. 29, 2015. doi: 10.1016/j.gene. 2015.06. 060. McCOOKE J.K. The mitochondrial genome of a Texas outbreak strain of the cattle tick, Rhipicephalus (Boophilus) microplus, derived from whole genome sequencing Pacific Biosciences and Illumina reads. Gene 571 135 141 2015 Available from: http://www.ncbi.nlm.nih.gov/pubmed/26117172 Oct. 29, 2015 10.1016/j.gene. 2015.06. 060 MEEUS, T. et al. Swift sympatric adaptation of a species of cattle tick to a new deer host in New Caledonia., Infection Genetics and Evolution v.10, p.976-983, 2010. Available from: <Available from: http://www.sciencedirect.com/science/article/pii/S1567134810001693 >. Accessed: Jan. 19, 2015. doi: 10.1016/j.meegid.2010.06.005. MEEUS T. Swift sympatric adaptation of a species of cattle tick to a new deer host in New Caledonia Infection Genetics and Evolution 10 976 983 2010 Available from: http://www.sciencedirect.com/science/article/pii/S1567134810001693 Jan. 19, 2015 10.1016/j.meegid.2010.06.005. MINNING, W. Beitrage zur Systematik und Morphologie der Zeckengattung Boophilus curtice. Z. Zeitschrift für Parasitenkunde, v.7, p.1-43, 1934. MINNING W. Beitrage zur Systematik und Morphologie der Zeckengattung Boophilus curtice. Z. Zeitschrift für Parasitenkunde 7 1 43 1934 MOHLER, J.R. Texas or tick fever and its prevention. Farm Bulletin of United States Department of Agriculture, v.238, p.1-44, 1906. MOHLER J.R. Texas or tick fever and its prevention Farm Bulletin of United States Department of Agriculture 238 1 44 1906 MURRELL, A. et al. A total-evidence phylogeny of ticks provides insights into the evolution of life cycles and biogeography., Molecular Phylogenetics and Evolution v.21, p.244-258, 2001. Available from: <Available from: http://www.sciencedirect.com/science/article/pii/S1055790301910188 >. Accessed: Jul. 23, 2014. doi: 10.10 06/mpev.2001.1018. MURRELL A. A total-evidence phylogeny of ticks provides insights into the evolution of life cycles and biogeography. Molecular Phylogenetics and Evolution 21 244 258 2001 Available from: http://www.sciencedirect.com/science/article/pii/S1055790301910188 Jul. 23, 2014 10.10 06/mpev.2001.1018 MURRELL, A. et al. Phylogenetic analyses of the rhipicephaline ticks indicate that the genus Rhipicephalus is paraphyletic., Molecular Phylogenetics and Evolution v.16, p.1-7, 2000. Available from: <Available from: http://www.sciencedirect.com/science/article/pii/S1055790300907620 >. Accessed: Sept. 27, 2014. doi: 10.1006/mpev.2000.0762. MURRELL A. Phylogenetic analyses of the rhipicephaline ticks indicate that the genus Rhipicephalus is paraphyletic. Molecular Phylogenetics and Evolution 16 1 7 2000 Available from: http://www.sciencedirect.com/science/article/pii/S1055790300907620 Sept. 27, 2014 10.1006/mpev.2000.0762. MURRELL, A. et al. The value of idiosyncratic markers and changes to conserved tRNA sequences from the mitochondrial genome of hard ticks (Acari: Ixodida: Ixodidae) for phylogenetic inference. Systematic Biology, v.52, p.296-310, 2003. Available from: <Available from: http://sysbio.oxfordournals.org/content/52/3/296.long >. Accessed: Jan. 12, 2015. doi: 10.1080/10635150390196957. MURRELL A. The value of idiosyncratic markers and changes to conserved tRNA sequences from the mitochondrial genome of hard ticks (Acari: Ixodida: Ixodidae) for phylogenetic inference. Systematic Biology 52 296 310 2003 Available from: http://sysbio.oxfordournals.org/content/52/3/296.long Jan. 12, 2015 10.1080/10635150390196957. MURRELL, A.; BARKER, S.C. Synonymy of Boophilus curtice, 1891 with Rhipicephalus koch, 1844 (Acari: Ixodidae). Systematic, Parasitology v.56, p.169-172, 2003. Available from: <Available from: http://www.ncbi.nlm.nih.gov/pubmed/14707501 >. Accessed: May 12, 2014. doi: 10.1023/B:SYPA.0000003802.36517.a0. MURRELL A. BARKER S.C. Synonymy of Boophilus curtice, 1891 with Rhipicephalus koch, 1844 (Acari: Ixodidae). Systematic, Parasitology 56 169 172 2003 Available from: http://www.ncbi.nlm.nih.gov/pubmed/14707501 May 12, 2014 10.1023/B:SYPA.0000003802.36517.a0. NEUMANN, L.G. Revision de la famille des Ixodides Mem. Bulletin de la Société zoologique de France, v.14, p.249-372, 1901. NEUMANN L.G. Revision de la famille des Ixodides Mem Bulletin de la Société zoologique de France 14 249 372 1901 OLWOCH J.M. et al. Climate change and the genus Rhipicephalus (Acari: Ixodidae) in Africa. Onderstepoort Journal of Veterinary Research, v.74, p.45-72, 2007. Available from: <Available from: http://www.ncbi.nlm.nih.gov/pubmed/17708153 >. Accessed: Sept. 21, 2014. doi: 10.4102/ojvr.v74i1.139. OLWOCH J.M. Climate change and the genus Rhipicephalus (Acari: Ixodidae) in Africa. Onderstepoort Journal of Veterinary Research 74 45 72 2007 Available from: http://www.ncbi.nlm.nih.gov/pubmed/17708153 Sept. 21, 2014 10.4102/ojvr.v74i1.139. PIESMAN, J.; EISEN, L. Prevention of tick-borne diseases. Annual Reviews of Entomology, v.53, p.323-343, 2008. Available from: <Available from: http://www.annualreviews.org >. Accessed: Dec. 12, 2014. doi: 10.1146/annurev.ento.53.103106.093429. PIESMAN J. EISEN L. Prevention of tick-borne diseases. Annual Reviews of Entomology 53 323 343 2008 Available from: http://www.annualreviews.org Dec. 12, 2014 10.1146/annurev.ento.53.103106.093429. POUND, J.M. et al. Evidence for role of white-tailed deer (Artiodactyla: Cervidae) in epizootiology of cattle ticks and southern cattle ticks (Acari: Ixodidae) in reinfestations along the Texas/Mexico border in south Texas: a review and update. Journal of Economic Entomology, v.103, p.211-218, 2010. Available from: <Available from: http://jee.oxfordjournals.org/ >. Accessed: Sept. 27, 2015. doi: 10.1603/EC09359. POUND J.M. Evidence for role of white-tailed deer (Artiodactyla: Cervidae) in epizootiology of cattle ticks and southern cattle ticks (Acari: Ixodidae) in reinfestations along the Texas/Mexico border in south Texas: a review and update. Journal of Economic Entomology 103 211 218 2010 Available from: http://jee.oxfordjournals.org/ Sept. 27, 2015 10.1603/EC09359. RECK, J. et al. First report of fluazuron resistance in Rhipicephalus microplus: a field tick population resistant to six classes of acaricides. Veterinary, Parasitology v.20, p.128-136, 2014. Available from: <Available from: http://www.ncbi.nlm.nih.gov/pubmed/24560364 >. Accessed: May 23, 2015. doi: 10.1016/j.vetpar.2014.01.012. RECK J. First report of fluazuron resistance in Rhipicephalus microplus: a field tick population resistant to six classes of acaricides. Veterinary, Parasitology 20 128 136 2014 Available from: http://www.ncbi.nlm.nih.gov/pubmed/24560364 May 23, 2015 10.1016/j.vetpar.2014.01.012. ROBERTS, F.H.S. The taxonomic status of the species of the genera Rhipicephalus koch and Boophilus curtice (Acarina: Ixodidae) occurring in Australia. Australian Journal of Zoology, v.13, p.491-523, 1965. Available from: <Available from: http://www.publish.csiro.au/paper/ZO9650491.htm >. Accessed: Dec. 05, 2014. doi: 10.1071/ZO9650491. ROBERTS F.H.S. The taxonomic status of the species of the genera Rhipicephalus koch and Boophilus curtice (Acarina: Ixodidae) occurring in Australia Australian Journal of Zoology 13 491 523 1965 Available from: http://www.publish.csiro.au/paper/ZO9650491.htm Dec. 05, 2014 10.1071/ZO9650491. SALMON, D.E.; STILES, C.W. The cattle ticks (Ixodoidea) of the United States. In: ANNUAL REPORT OF THE UNITED STATE, 17, 1901, Washington: Department of Agriculture, Bureau of Animal Industry, 1901. p.380-491. SALMON D.E. STILES C.W. The cattle ticks (Ixodoidea) of the United States. ANNUAL REPORT OF THE UNITED STATE, 17 1901 Washington Department of Agriculture, Bureau of Animal Industry 1901 380 491 SATTLER, P.W. et al. Genetic similarity and variability between natural populations and laboratory colonies of North American Boophilus (Acari: Ixodidae). Journal of, Parasitology v.72, p.95-100, 1986. Available from: <Available from: http://www.ncbi.nlm.nih.gov/pubmed/3712179 >. Accessed: Dec. 12, 2014. doi: 10.2307 /3281799. SATTLER P.W. Genetic similarity and variability between natural populations and laboratory colonies of North American Boophilus (Acari: Ixodidae). Journal of, Parasitology 72 95 100 1986 Available from: http://www.ncbi.nlm.nih.gov/pubmed/3712179 Dec. 12, 2014 10.2307 /3281799 SAUER, J.R. et al. Salivary glands in ixodid ticks: control and mechanism of secretion. Journal of Insect Physiology, v.46, p.10691078, 2000. Available from: <Available from: http://www.sciencedirect.com/science/article/pii/S0022191099002103 >. Accessed: Dec. 23, 2014. doi: 10.1016/S0022-1910(99)00210-3. SAUER J.R. Salivary glands in ixodid ticks: control and mechanism of secretion. Journal of Insect Physiology 46 10691078 10691078 2000 Available from: http://www.sciencedirect.com/science/article/pii/S0022191099002103 Dec. 23, 2014 10.1016/S0022-1910(99)00210-3. SPICKETT, A.M.; MALAN, J.R. Genetic incompatibility between Boophilus decoloratus (Koch, 1844) and Boophilus microplus (Canestrini, 1888) and hybrid sterility of Australian and South African Boophilus microplus (Acarina: Ixodidae)., Onderstepoort Journal of Veterinary Research v.45, p.149-153, 1978. SPICKETT A.M. MALAN J.R. Genetic incompatibility between Boophilus decoloratus (Koch, 1844) and Boophilus microplus (Canestrini, 1888) and hybrid sterility of Australian and South African Boophilus microplus (Acarina: Ixodidae) Onderstepoort Journal of Veterinary Research 45 149 153 1978 THOMPSON, G.D. et al. Hybrid sterility in cattle ticks. Experientia, v.37, p.127-128, 1981. Available from: <Available from: http://www.ncbi.nlm.nih.gov/pubmed/571591 >. Accessed: Dec. 21, 2014. doi: 10.1007/BF01963186. THOMPSON G.D. Hybrid sterility in cattle ticks. Experientia 37 127 128 1981 Available from: http://www.ncbi.nlm.nih.gov/pubmed/571591 Dec. 21, 2014 10.1007/BF01963186. TONNESEN, M.H. et al. Displacement of Boophilus decoloratus by Boophilus microplus in the Soutpansberg region, Limpopo province, South Africa., Experimental and Applied Acarology v.32, p.199-208, 2004. Available from: <Available from: http://www.ncbi.nlm.nih.gov/pubmed/15139085 >. Accessed: Jan. 15, 2015. doi: 10.1023/B:APPA.0000021789.44411.b5. TONNESEN M.H. Displacement of Boophilus decoloratus by Boophilus microplus in the Soutpansberg region, Limpopo province, South Africa Experimental and Applied Acarology 32 199 208 2004 Available from: http://www.ncbi.nlm.nih.gov/pubmed/15139085 Jan. 15, 2015 10.1023/B:APPA.0000021789.44411.b5 UILENBERG, G. Boophilus (Uroboophilus) fallax Minning, 1934 synonyme de Boophilus microplus (Canestrini, 1887) (Ixodidae). Revue d'elevage et de Medecine Veterinaire des Pays Tropicaux, v.15, p.387-398,1962. UILENBERG G. (Uroboophilus) fallax Minning, 1934 synonyme de Boophilus microplus (Canestrini, 1887) (Ixodidae). Revue d'elevage et de Medecine Veterinaire des Pays Tropicaux 15 387 398 1962 VERGES J. Les tiques du Betail. Methodes d'eradication. Noumea: Imprimeries reunies, 1944. 72p. VERGES J. Les tiques du Betail Methodes d'eradication. Noumea Imprimeries reunies 1944 72 WALKER, A.R. et al. Ticks of domestic animals in Africa: a guide to identification of species. Bioscience Reports, Edinburgh, 2003. 221p. WALKER A.R. Ticks of domestic animals in Africa: a guide to identification of species. Bioscience Reports 2003 221 WILLADSEN, P. Anti-tick vaccines., Parasitology v.129, p.S367-S387, 2004. Available from: <Available from: http://www.ncbi.nlm.nih.gov/pubmed/15938519 >. Accessed: Dec. 07, 2014. doi: 10.1017/S003118200300 4657. WILLADSEN P. Anti-tick vaccines. Parasitology 129 S367 S387 2004 Available from: http://www.ncbi.nlm.nih.gov/pubmed/15938519 Dec. 07, 2014 10.1017/S003118200300 4657 XU, Z. et al. Complete mitochondrial genome of Rhipicephalus simus. Mitochondrial DNA: The Journal of DNA Mapping, Sequencing, and Analysis. p 888-889. Accessed from: <http:// www.tandfonline.com/>. Accessed: Feb. 10, 2016. doi: 10.3109/19401736. 2014.919490. XU Z. Complete mitochondrial genome of Rhipicephalus simus. Mitochondrial DNA: The Journal of DNA Mapping, Sequencing, and Analysis. 888 889 http:// www.tandfonline.com/ 10 02 2016 10.3109/19401736. 2014.919490 1 CR-2015-1416