Ticks play a crucial role in ecosystem dynamics and disease transmission, necessitating accurate species identification for epidemiological and ecological research. South Africa, despite occupying only 0.82 % of the world's land area, harbors approximately 11 % of global tick species richness. This study provides an updated, authoritative checklist of South African tick species curated by the National Tick Collection of South Africa (NTCSA), housed at the Agricultural Research Council - Onderstepoort Veterinary Research (ARCOVR). Previous lists for South Africa presented outdated taxonomic classifications and incomplete data on taxonomic taxa. The NTCSA checklist aims to correct inaccuracies, incorporating the most recent literature, molecular systematics, and expert curation. A total of 110 tick species across three families (Ixodidae, Argasidae, Nuttalliellidae) are documented, with corrections to genera classifications and species distributions indicated. This list serves as a reference for researchers, veterinarians, and policymakers involved in tick diversity, vector control, and conservation efforts. Continuous updates integrating morphological and molecular assessments will further enhance our understanding of tick diversity and distribution in South Africa. The wide extralimital distribution of various species extend the taxonomic observations to the African continent, while the general taxonomic observations presented summarize our understanding of tick taxonomy at global level.

The Rhipicephalus sanguineus group, the brown dog ticks, are cosmopolitan and doubtless the most important ticks of domestic dogs, clinically and economically. Despite four decades of taxonomic enquiry with nucleotide sequences and morphology, the taxonomy of the R. sanguineus group is confused, even chaotic. We provide 13 new mitochondrial (mt) genomes and internal transcribed spacer 2 (ITS2) sequences from nine localities in Australia, Israel and Japan. We inferred phylogenetic trees from 10 mt protein-coding genes (9,514 bp), as well as partial cox1, ITS2, 12S, and 16S rRNA genes, to resolve to common clades the >2,000 nucleotide sequences in GenBank from the R. sanguineus group. Then we applied three species delimitation protocols to 60 entire mt genomes (ca. 15,000 bp) and 162 partial cox1 sequences (472 bp): Automatic Barcode Gap discovery, Assemble Species by Automatic Partitioning, and Poisson Tree Process. We considered pairwise genetic differences and Tamura-Nei genetic distances among 60 entire mt genomes and 162 partial cox1 sequences. We found 16 species-level clades (clades A to P) that we hypothesise represent at least 16 species in the R. sanguineus group. These clades had intra-clade differences of <3.8% (entire mt genomes) and <5.1% (partial cox1) whereas the inter-clade differences were >7.7% (entire mt genomes) and >4.5% (partial cox1). We assigned the species names Rhipicephalus linnaei (Audouin, 1826), Rhipicephalus rutilus (Koch, 1844), Rhipicephalus secundus (Feldman-Muhsam, 1952) and R. sanguineus (Latreille, 1806) to clades A, C, D, and K, respectively. And we hypothesise that the names Rhipicephalus camicasi (Morel, Mouchet & Rodhain, 1976), Rhipicephalus turanicus (Pomerantsev, 1940), Rhipicephalus guilhoni (Morel & Vassilades, 1963), Rhipicephalus sulcatus (Neumann, 1908), Rhipicephalus rossicus (Yakimov & Kol-Yakimova, 1911), Rhipicephalus pumilio (Schulze, 1935) and Rhipicephalus pusillus (Gil Collado, 1936) apply to clades B, E, H, J, M, N and O, respectively. The newly described Rhipicephalus hibericus (Millán, Rodriguez-Pastor & Estrada-Peña, 2024) was genetically indistinguishable from R. sanguineus in clade K and thus is a synonym of R. sanguineus. We could not assign names to clades F (USA, Hungary), I (India, Pakistan), L (Nigeria), G (China, Kazakhstan), and P (Cameroon): some or all of these five clades may be new species in the R. sanguineus group. Our haplotype network of partial mt genes (cox1, cytb and nad2) revealed much genetic similarity among geographically distant populations of R. linnaei. This indicates recent dispersal, likely originating in Africa or the Middle East, since African populations were more genetically diverse than populations in other parts of the world.

Background The presence of alpha-gal in tick salivary antigens has been implicated to initiate host IgE responses resulting in sensitization to alpha-gal. Objectives We investigated the presence of alpha-gal in different anatomic locations in native South African tick species at different feeding and developmental stages and the ability of sera from our allergic cohort to bind to these tick proteins. Methods Alpha-gal containing proteins in lab reared ticks at different feeding and developmental stages were detected through western blotting and immunohistochemical staining. IgE and IgG4 towards the tick proteins was analysed via ELISA. Results There is differential expression of alpha-gal in endemic South African ticks Amblyomma hebraeum and Rhipicephalus evertsi. Immunoblotting demonstrated an increase in the prominence of alpha-gal containing protein bands in salivary glands to be proportional to feeding time. Alpha-gal in both fed and unfed ticks was localized in the salivary acini and testes. IgE and IgG4 to A. hebraeum antigens was significantly raised in alpha-gal allergic individuals. There was a correlation between anti-A. hebraeum IgE and anti-alpha-gal IgE in the alpha-gal allergic group. Inhibition of human serum anti-alpha-gal IgE to A. hebraeum and R. evertsi proteins was significantly reduced by the addition of bovine thyroglobulin. Conclusion We anticipate repeated exposure to differentially expressed A. hebraeum and R. evertsi alpha-gal-containing salivary proteins during feeding to cause alpha-gal sensitization.

The identity of Ixodes (Afrixodes) ugandanus Neumann, 1906 (Acari: Ixodidae) is established and its male and female are redescribed based on specimens collected on rodents (Rodentia: Muridae, Thryonomyidae) in Ethiopia and Uganda. Lectotype of I. ugandanus is designated here. Ixodes (Afrixodes) ampullaceus Warburton, 1933 is reestablished as a valid species and its female is redescribed based on specimens collected on rodents (Rodentia: Thryonomyidae), carnivorans (Carnivora: Viverridae) and a bird (Galliformes: Numididae) in Kenya, Malawi, Mozambique, South Africa, Tanzania, and Zimbabwe. Ixodes mossambicensis Santos Dias, 1952 is synonymized with I. ampullaceus. Male and female of Ixodes (Afrixodes) aulacodi Arthur, 1956 are redescribed in greater detail based on specimens collected on rodents (Rodentia: Thryonomyidae) in Benin, Cameroon, and Ivory Coast. All available literature data on these taxa are critically analyzed and discussed. We sequenced the mitochondrial genome of I. aulacodi, compared it to other Ixodes Latreille, 1795 species, and according to our phylogeny from 10 protein-coding mitochondrial genes of 33 Ixodes species, I. aulacodi forms a clade with the only other Afrixodes Morel, 1966 having a sequenced complete mitochondrial genome, Ixodes rubicundus Neumann, 1904.

Background:The presence of galactose-α-1,3-galactose (alpha-gal) in tick salivary antigens has been implicated to initiate host IgE responses resulting in sensitization to alpha-gal. Objective:We sought to investigate the presence of alpha-gal in different anatomic locations in native South African tick species at different feeding and developmental stages and the ability of sera from our allergic cohort to bind to these tick proteins. Methods:Alpha-gal-containing proteins in laboratory-reared ticks at different feeding and developmental stages were detected through Western blotting and immunohistochemical staining. IgE and IgG4 toward the tick proteins were analyzed via ELISA. Results:There is differential expression of alpha-gal in endemic South African ticks Amblyomma hebraeum and Rhipicephalus evertsi. Immunoblotting demonstrated an increase in the prominence of alpha-gal-containing protein bands in salivary glands to be proportional to feeding time. Alpha-gal in both fed and unfed ticks was localized in the salivary acini and testes. IgE and IgG4 to A hebraeum antigens were significantly raised in alpha-gal-allergic individuals. There was a correlation between anti-A hebraeum IgE and anti-alpha-gal IgE in the alpha-gal-allergic group. Inhibition of human serum anti-alpha-gal IgE to A hebraeum and R evertsi proteins was significantly reduced by the addition of bovine thyroglobulin. Conclusions:We anticipate repeated exposure to differentially expressed A hebraeum and R evertsi alpha-gal-containing salivary proteins during feeding to cause alpha-gal sensitization.

Mitochondrial (mt) genomes have played a major role in elucidating evolutionary relationships in ticks (Ixodida), especially in soft ticks (Argasidae). Mitochondrial genomes have been sequenced for representatives of most of the genera and other major lineages. This includes members of the Afrotropical, Nearctic, Neotropical and Palearctic Pavlovskyella Pospelova-Shtrom, 1950, which is a subgenus of Ornithodoros Koch, 1844. These continent-associated lineages do not form a monophyletic group: rather, the subgenus Pavlovskyella is paraphyletic. The only zoogeographic region for which mt genomes are not available is the Australasian region. Here we report the first mt genome of an Australasian Pavlovskyella: Ornithodoros (Pavlovskyella) gurneyi Warburton, 1926, the kangaroo soft tick. [The companion paper to the present work, Barker et al. (2025), presents the mt genome, 18S and 28S rRNA of the only other known Australasian Pavlovskyella, McMillan’s Australian tree-hollow argasid, Ornithodoros (Pavlovskyella) macmillani Hoogstraal & Kohls, 1966.] Our phylogenetic trees reveal that the two Australasian Pavlovskyella have their own clade (i.e. are sister-taxa) which does not have a sister-group relationship with species of Pavlovskyella from any of the four other zoogeographic regions. We propose that the Pavlovskyella from the Afrotropical, Australasian, Nearctic, Neotropical and Palearctic zoogeographic regions have independent evolutionary histories and thus are best considered different genera. Molecular dating leads us to propose that continental drift may have caused the evolution of five different lineages of Pavlovskyella, each in a different zoogeographic region of the world.

The genus Amblyomma contains the highest percentage of reptile-associated ticks, and comprises approximately nine subgenera. One of these subgenera is Adenopleura, which also encompasses Amblyomma javanense, and its type species Amblyomma compressum. This study describes a new Amblyomma species associated with Bengal monitor lizards (Varanus bengalensis) based on morphology and its mitogenome in Khyber Pakhtunkhwa, Pakistan. Reptiles belonging to different genera were examined for Amblyomma ticks and only the monitor lizard was infested with ticks in the District Bajaur. Collected Amblyomma cf. javanense ticks were analyzed and formally described as a new species. Overall, 57 A. cf. javanense ticks were collected on monitor lizards (4/27) with a 15% prevalence of infestation, 2.1 mean abundance, and 14.3 mean intensity. Ticks comprised males (n = 23, 40%), females (n = 14, 25%) and nymphs (n = 20, 35%), while no larvae were found. BLAST analysis of A. cf. javanense sequences showed the following maximum identities; 98.25% with undetermined Amblyomma species based on 12S rRNA, 96.07% with A. javanense based on 16S rRNA, 99.56% and 90.95% with an Amblyomma sp. and A. javanense, respectively, based on ITS2. Moreover, the mitochondrial genome of A. cf. javanense showed maximum identities of 80.75%, 80.48% and 79.42% with Amblyomma testudinarium, A. javanense, and Amblyomma sp., respectively. The phylogenetic analysis of A. cf. javanense revealed that its 12S rRNA and 16S rRNA are closely related to an Amblyomma sp. and A. javanense, respectively, from Sri Lanka, its ITS2 is closely related to A. javanense from China and an Amblyomma sp. from Sri Lanka, and its mitogenome is closely related to A. javanense and Amblyomma sp. from China. The pairwise distance analysis resulted in divergence of 0-1.71% (12S rRNA), 0-17.5% (16S rRNA), 0-9.1% (ITS2) and 0-20.5% (mitochondrial genome). We also contributed the full-length mitochondrial genome sequence of A. compressum and showed that this species does not share a most recent common ancestor with A. javanense. As the subgenus Adenopleura is paraphyletic, this study could help to understand the systematics and phylogeny of this taxon.

The genus Ornithodoros is notably diverse within the family Argasidae, comprising approximately 134 species distributed among 4 subgenera, 1 of which is the subgenus Pavlovskyella. In an earlier study, we identified distinct soft ticks as Ornithodoros (Pavlovskyella) sp., which were collected from animal shelters in Khyber Pakhtunkhwa, Pakistan. Providing additional collections from that same locality and a comprehensive analysis involving detailed morphological and mitogenome-based comparisons with closely related species, this study formally designates a novel species for these specimens. Adults and late-instar nymphs of the new species display a dorsoventral groove, small cheeks not covering the capitulum, 5 small even humps on tarsus I and a transverse postanal groove intersecting the median postanal groove perpendicularly. It also lacks a tuft of setae on the ventral surface of the hood which separates the novel species from Ornithodoros papillipes. Ventral chaetotaxy of tarsus IV indicates 4–7 setal pairs in nymphs and 5–7 pairs in adults that separate the new species from Ornithodoros tholozani sensu stricto and Ornithodoros crossi, 2 morphologically closely related species that occur in geographical proximity. Phylogenetic analyses of the full-length mitochondrial genome and the 18S and 28S ribosomal RNA genes, combined with pairwise nucleotide comparisons of cox1, cox2, atp8, atp6, cox3, nad3, nad5, nad4, nad4L, nad6, cytb, nad1, nad2, 12S rDNA, 16S rDNA, 18S rDNA and 28S rDNA further support that the new species belongs to the Pavlovskyella subgenus, clustering with O. tholozani, Ornithodoros verrucosus and Ornithodoros tartakovskyi.

We describe a new genus Cryptocroton n. gen. for Amblyomma papuanum Hirst, 1914, a tick of North Queensland, Australia, and Papua New Guinea.

Ixodid ticks are obligate blood-feeding arthropods and important vectors of pathogens. In Mallorca, almost no data on the tick fauna are available. Herein, we investigated ticks and tick-borne pathogens in ticks collected from dogs, a cat and humans in Mallorca as result of a citizen science project. A total of 91 ticks were received from German tourists and residents in Mallorca. Ticks were collected from March to October 2023 from dogs, cat and humans, morphologically and genetically identified and tested for pathogens by PCRs. Six tick species could be identified: Ixodes ricinus (n = 2), Ixodes ventalloi (n = 1), Hyalomma lusitanicum (n = 7), Hyalomma marginatum (n = 1), Rhipicephalus sanguineus s.l. (n = 71) and Rhipicephalus pusillus (n = 9). Rhipicephalus sanguineus s.l. adults were collected from dogs and four females from a cat and the 16S rDNA sequences identified it as Rh. sanguineus s.s. Hyalomma lusitanicum was collected from 1 human, 1 dog and 5 specimens were collected from the ground in the community of Santanyi, together with one H. marginatum male. This is the first report of Hyalomma marginatum in Mallorca. Both I. ricinus were collected from humans and I. ventalloi female was collected from a dog. All ticks tested negative for Anaplasma phagocytophilum, Coxiella spp., Francisella spp., and piroplasms. In 32/71 (45%) specimens of Rh. sanguineus s.s., Rickettsia spp. could be detected and in 18/32 (56.2%) sequenced tick DNAs R. massiliae was identified. Ixodes ventalloi female and both I. ricinus tested positive in the screening PCR, but the sequencing for the identification of the Rickettsia sp. failed.

Two major families exist in ticks, the Argasidae and Ixodidae. The Argasidae comprise 2 sub-families, Argasinae and Ornithodorinae. The placement into subfamilies illuminate differences in morphological and molecular systematics and is important since it provides insight into evolutionary divergence within this family. It also identifies fundamental gaps in our understanding of argasid evolution that provide directions for future research. Molecular systematics based on mitochondrial genomics and 18S/28S ribosomal RNA confirmed the placement of various genera and subgenera into the Argasinae: Argas (including Argas and Persicargas ), Navis , Ogadenus , Otobius lagophilus , Proknekalia , Secretargas and the Ornithodorinae: Alectorobius , Antricola (including Antricola and Parantricola ), Carios , Chiropterargas , Nothoaspis , Ornithodoros (including Microargas , Ornamentum , Ornithodoros sensu strictu, Pavlovskyella ), Otobius sensu strictu, Reticulinasus and Subparmatus . The position of Alveonasus remains controversial since traditional taxonomy placed it in the Ornithodorinae, while cladistic and limited molecular analysis placed it in the Argasinae. The current study aimed to resolve the systematic position of Alveonasus using mitochondrial genomic and 18S/28S ribosomal RNA systematics by sequencing the type species Alveonasus lahorensis from Pakistan. In addition, the mitochondrial genomes for Argas reflexus and Alectorobius kelleyi are reported from Germany and the USA, respectively. The systematic data unambiguously place Alveonasus in the Argasinae and also suggest that Alveonasus may be another paraphyletic genus.

Ticks are composed of 3 extant families (Argasidae, Ixodidae and Nuttalliellidae) and 2 extinct families (Deinocrotonidae and Khimairidae). The Nuttalliellidae possess one extant species ( Nuttalliella namaqua ) limited to the Afrotropic region. A basal relationship to the hard and soft tick families and its limited distribution suggested an origin for ticks in the Afrotropics. The Deinocrotonidae has been found in Burmese amber from Myanmar and Iberian amber from Spain, suggesting a wider distribution of the lineage composed of Deinocrotonidae and Nuttalliellidae. The current study describes 8 fossils from mid-Cretaceous (ca. 100 Ma) Burmese amber: 2 Deinocroton species ( Deinocroton bicornis sp. nov.; Deinocroton lacrimus sp. nov.), 5 Nuttalliella species ( Nuttalliella gratae sp. nov., Nuttalliella tuberculata sp. nov., Nuttalliella placaventrala sp. nov., Nuttalliella odyssea sp. nov., Nuttalliella tropicasylvae sp. nov.) and a new genus and species ( Legionaris nov. gen., Legionaris robustus sp. nov.). The argument is advanced that Deinocroton do not warrant its own family, but forms part of the Nuttalliellidae comprising 3 genera, Deinocroton , Legionaris nov. gen. and Nuttalliella ). Affinities of Burmese tick fossils to the Australasian region, specifically related to rifting of the Burma terrane from northern Australia ~150 million years ago, suggest that Nuttalliella had a much wider distribution than its current limited distribution. The distribution of Nuttalliella likely stretched from Africa over Antarctica and much of Australia, suggesting that extant members of this family may still be found in Australia. Considerations for the geographic origins of ticks conclude that an Afrotropic origin can as yet not be discarded.

We used entire mitochondrial (mt) genome sequences (14.5-15 kbp) to resolve the phylogeny of the four main lineages of the Haematobothrion ticks: Alloceraea, Archaeocroton, Bothriocroton and Haemaphysalis. In our phylogenetic trees, Alloceraea was the sister to Archaeocroton sphenodonti, a tick of an archetypal reptile, the tuatara, from New Zealand, to the exclusion of the rest of the species of Haemaphysalis. The mt genomes of all four of the Alloceraea species that have been sequenced so far had a substantial insert, 132-312 bp, between the tRNA-Glu (E) gene and the nad1 gene in their mt genomes. This insert was not found in any of the other eight subgenera of Haemaphysalis. The mt genomes of 13 species of Haemaphysalis from NCBI GenBank were added to the most recent data set on Haemaphysalis and its close relatives to help resolve the phylogeny of Haemaphysalis, including five new subgenera of Haemaphysalis not previously considered by other authors: Allophysalis (structurally primitive), Aboimisalis (structurally primitive), Herpetobia (structurally intermediate), Ornithophysalis (structurally advanced) and Segalia (structurally advanced). We elevated Alloceraea Schulze, 1919 to the status of genus because Alloceraea Schulze, 1919 is phylogenetically distinct from the other subgenera of Haemaphysalis. Moreover, we propose that the subgenus Allophysalis is the sister to the rest of the Haemaphysalis (14 subgenera) and that the 'structurally primitive' subgenera Hoogstraal and Kim comprise early diverging lineages. Our matrices of the pairwise genetic difference (percent) of mt genomes and partial 16S rRNA sequences indicated that the mt genome sequence of Al. kitaokai (gb# OM368280) may not be Al. kitaokai Hoogstraal, 1969 but rather another species of Alloceraea. In a similar way, the mt genome sequence of H. (Herpetobia) nepalensis Hoogstraal, 1962 (gb# NC_064124) was only 2% genetically different to that of H. (Allophysalis) tibetensis Hoogstraal, 1965 (gb# OM368293): this indicates to us that they are the same species. Alloceraea cretacea may be better placed in a genus other than Alloceraea Schulze, 1919. Reptiles may have been the host to the most recent common ancestor of Archaeocroton and Alloceraea.

Theileria parva are intracellular protozoal parasites responsible for three disease syndromes in cattle, namely East Coast fever (ECF), Corridor disease (CD) and Zimbabwean theileriosis. The increase in reports of CD outbreaks in recent years has raised questions about the probability of adaptation of buffalo-derived T. parva strains in cattle herds adjacent to game reserves. A cross-sectional study was conducted from March 2016 to December 2018 to investigate the extent of occurrence of T. parva infections in cattle in the CD-controlled area of KwaZulu-Natal Province. Blood samples were collected from 1137 cattle from 14 herds and analysed by quantitative real-time PCR (qPCR) and indirect fluorescent antibody test (IFAT) to determine the prevalence of T. parva. A total of 484 samples from 4 of the 14 herds were further tested on qPCR for the presence of T. taurotragi infections. The data were analysed using descriptive statistics and a chi-square test was used to assess association between variables. The overall prevalence of T. parva was 1.3% (95%CI:1-2%) and 19.9% (95%CI:17-22%) on qPCR and IFAT, respectively. The qPCR positive samples were detected in March and May while IFAT positive samples were detected in all seasons sampled, with higher numbers during summer months. The Pearson Chi-squared test showed that T. parva prevalence rates based on both qPCR and IFAT were positively associated with herds with previous history of CD outbreaks (chi(2) = 8.594, p = 0.003; chi(2) = 69.513, p < 0.001, respectively). The overall prevalence of T. taurotragi was 39.4% (95% CI: 35-44%) with the herd-level prevalence ranging between 35.0% and 43.4%. Possible cross-reaction of T. parva IFAT to T. taurotragi was detected on few samples, however, there was no significant association between T. taurotragi infections and IFAT positivity (chi(2) = 0.829, p = 0.363). Results from this study demonstrated the extent of occurrence of subclinical carriers and the level of exposure to T. parva infections in cattle populations at a livestock/game interface area of KwaZulu-Natal Province. The molecular and seroprevalence rates were low when compared with other areas where cattle-adapted T. parva infections are endemic. The adaptation of buffalo-derived T. parva in cattle population resulting in cattle-cattle transmissions seem to be unlikely under the current epidemiological state.

Eleonora's falcon (Falco eleonorae Géné, 1839) is a well-known long-distance migrant of the Afro-Palaearctic flyway, a summer breeder of the Mediterranean region and North-west Africa and a winter resident of Madagascar and surrounding areas, thus characterized as a double endemic. Within the context of a long-term monitoring and conservation programme on Antikythira Island, Greece, which accommodates one of the largest concentrations of breeding pairs of Eleonora's falcons globally, birds were subjected to regular inspections for the presence of ticks from 2017 to 2023. In total, 104 adults and 149 nymphs (all belonging to Haemaphysalis genus) were collected. All ticks, apart from 2 nymphs, exhibited broadly salient palpi and did not possess the pronounced palpal segment 2 spurs or spur-like angles that are characteristic of adults, nymphs and most larvae of Rhipistoma, thus placed them in the Ornithophysalis subgenus. Following comprehensive morphological assessment and genetic analysis of the mitochondrial genome by means of next-generation sequencing of both adult and nymphal stages of the ticks, our empirical findings substantiate the delineation of a previously unclassified species. This taxonomic assignment situates the newly described species within the Ornithophysalis subgenus and the Haemaphysalis doenitzi group, marking its presence for the first time within the Western Palaearctic region.

Background Amblyomma is the third most diversified genus of Ixodidae that is distributed across the Indomalayan, Afrotropical, Australasian (IAA), Nearctic and Neotropical biogeographic ecoregions, reaching in the Neotropic its highest diversity. There have been hints in previously published phylogenetic trees from mitochondrial genome, nuclear rRNA, from combinations of both and morphology that the Australasian Amblyomma or the Australasian Amblyomma plus the Amblyomma species from the southern cone of South America, might be sister-group to the Amblyomma of the rest of the world. However, a stable phylogenetic framework of Amblyomma for a better understanding of the biogeographic patterns underpinning its diversification is lacking. Methods We used genomic techniques to sequence complete and nearly complete mitochondrial genomes –ca. 15 kbp– as well as the nuclear ribosomal cluster –ca. 8 kbp– for 17 Amblyomma ticks in order to study the phylogeny and biogeographic pattern of the genus Amblyomma , with particular emphasis on the Neotropical region. The new genomic information generated here together with genomic information available on 43 ticks (22 other Amblyomma species and 21 other hard ticks–as outgroup–) were used to perform probabilistic methods of phylogenetic and biogeographic inferences and time-tree estimation using biogeographic dates. Results In the present paper, we present the strongest evidence yet that Australasian Amblyomma may indeed be the sister-group to the Amblyomma of the rest of the world (species that occur mainly in the Neotropical and Afrotropical zoogeographic regions). Our results showed that all Amblyomma subgenera ( Cernyomma , Anastosiella , Xiphiastor , Adenopleura , Aponomma and Dermiomma ) are not monophyletic, except for Walkeriana and Amblyomma . Likewise, our best biogeographic scenario supports the origin of Amblyomma and its posterior diversification in the southern hemisphere at 47.8 and 36.8 Mya, respectively. This diversification could be associated with the end of the connection of Australasia and Neotropical ecoregions by the Antarctic land bridge. Also, the biogeographic analyses let us see the colonization patterns of some neotropical Amblyomma species to the Nearctic. Conclusions We found strong evidence that the main theater of diversification of Amblyomma was the southern hemisphere, potentially driven by the Antarctic Bridge's intermittent connection in the late Eocene. In addition, the subgeneric classification of Amblyomma lacks evolutionary support. Future studies using denser taxonomic sampling may lead to new findings on the phylogenetic relationships and biogeographic history of Amblyomma genus. Graphical Abstract

The study of tick evolution may be classified into disciplines such as taxonomy and systematics, biogeography, evolution and development (evo-devo), ecology, and hematophagy. These disciplines overlap and impact each other to various extents. Advances in one field may lead to paradigm shifts in our understanding of tick evolution not apparent to other fields. The current study considers paradigm shifts that occurred, are in the process, or may occur in future for the disciplines that study tick evolution. Some disciplines have undergone significant changes, while others may still be developing their own paradigms. Integration of these various disciplines is essential to come to a holistic view of tick evolution; however, maturation of paradigms may be necessary before this vision can be attained.