Effects of Tectonics and Large Scale Climatic Changes On the Evolutionary History of \u3ci\u3eHyalomma\u3c/i\u3e ticks

Hyalomma Koch, 1844 are ixodid ticks that infest mammals, birds and reptiles, to which 27 recognized species occur across the Afrotropical, Palearctic and Oriental regions. Despite their medical and veterinary importance, the evolutionary history of the group is enigmatic. To investigate various taxonomic hypotheses based on morphology, and also some of the mechanisms involved in the diversification of the genus, we sequenced and analysed data derived from two mtDNA fragments, three nuclear DNA genes and 47 morphological characters. Bayesian and Parsimony analyses based on the combined data (2242 characters for 84 taxa) provided maximum resolution and strongly supported the monophyly of Hyalomma and the subgenus Euhyalomma Filippova, 1984 (including H. punt Hoogstraal, Kaiser and Pedersen, 1969). A predicted close evolutionary association was found between morphologically similar H. dromedarii Koch, 1844, H. somalicum Tonelli Rondelli, 1935, H. impeltatum Schulze and Schlottke, 1929 and H. punt, and together they form a sister lineage to H. asiaticum Schulze and Schlottke, 1929, H. schulzei Olenev, 1931 and H. scupense Schulze, 1919. Congruent with morphological suggestions, H. anatolicum Koch, 1844, H. excavatum Koch, 1844 and H. lusitanicum Koch, 1844 form a clade and so also H. glabrum Delpy, 1949, H. marginatum Koch, 1844, H. turanicum Pomerantzev, 1946 and H. rufipes Koch, 1844. Wide scale continental sampling revealed cryptic divergences within African H. truncatum Koch, 1844 and H. rufipes and suggested that the taxonomy of these lineages is in need of a revision. The most basal lineages in Hyalomma represent taxa currently confined to Eurasia and molecular clock estimates suggest that members of the genus started to diverge approximately 36.25 million years ago (Mya). The early diversification event coincides well with the collision of the Indian and Eurasian Plates, an event that was also characterized by large scale faunal turnover in the region. Using S-Diva, we also propose that the closure of the Tethyan seaway allowed for the genus to first enter Africa approximately 17.73 Mya. In concert, our data supports the notion that tectonic events and large scale global changes in the environment contributed significantly to produce the rich species diversity currently found in the genus Hyalomma

FIGURE 10 in Description of eight new species of Ixodes Latreille, 1795 (Acari: Ixodidae) and redescription of I. auritulus Neumann, 1904, parasites of birds in the Australasian, Nearctic and Neotropical Regions

FIGURE 10. Map showing the known geographical distribution of Ixodes auritulus Neumann, 1904, I. brevisetosus n. sp., I. contrarius n. sp., I. guglielmonei n. sp., I. insulae n. sp., I. moralesi n. sp., I. rio n. sp., I. robbinsi n. sp. and I. tinamou n. sp. A, Filled star shows the type-locality and filled circles show confirmed localities of I. auritulus; B, Filled asterisk shows the type-locality and filled circles show confirmed localities of I. brevisetosus n. sp., filled star shows the type-locality of I. insulae n. sp.; C, Filled triangle shows the type-locality of I. contrarius n. sp., filled cross shows the type-locality and filled circle shows confirmed locality of I. moralesi n. sp.; D, Filled star shows the type-locality and filled circles show confirmed localities of I. guglielmonei n. sp.; E, Filled star shows the type-locality and filled circle shows confirmed locality of I. rio n. sp., filled asterisk shows the type-locality of I. tinamou n. sp.; F, Filled star shows the type-locality and filled circles show confirmed localities of I. robbinsi n. sp.Published as part of <i>Apanaskevich, Dmitry A., Apanaskevich, Maria A., Klimov, Pavel B., Edgy, Brett M., Bermúdez, Sergio E., Labruna, Marcelo B., Korzeev, Andrei I. & Barker, Stephen C., 2022, Description of eight new species of Ixodes Latreille, 1795 (Acari: Ixodidae) and redescription of I. auritulus Neumann, 1904, parasites of birds in the Australasian, Nearctic and Neotropical Regions, pp. 1-73 in Zootaxa 5173 (1)</i> on page 16, DOI: 10.11646/zootaxa.5173.1.1, <a href="http://zenodo.org/record/6966076">http://zenodo.org/record/6966076</a&gt

FIGURE 28. Ixodes rio n in Description of eight new species of Ixodes Latreille, 1795 (Acari: Ixodidae) and redescription of I. auritulus Neumann, 1904, parasites of birds in the Australasian, Nearctic and Neotropical Regions

FIGURE 28. Ixodes rio n. sp., scanning electron micrographs of female (Alto do Itatiaia, Serra do Itatiaia, Rio de Janeiro, Brazil, USNMENT 00987157, paratype). A, Idiosoma, dorsal view; B, Scutum, dorsal view; C, Scutum, dorsolateral view; D, Alloscutal setae in central area of alloscutum; E, Alloscutal setae in area of alloscutum medial to marginal grooves; F, Idiosoma, ventral view. Scale-bars: A, F, 0.5 mm; B, C, 0.2 mm; D, E, 0.05 mm. SEMs by D.A. ApanaskevichPublished as part of Apanaskevich, Dmitry A., Apanaskevich, Maria A., Klimov, Pavel B., Edgy, Brett M., Bermúdez, Sergio E., Labruna, Marcelo B., Korzeev, Andrei I. & Barker, Stephen C., 2022, Description of eight new species of Ixodes Latreille, 1795 (Acari: Ixodidae) and redescription of I. auritulus Neumann, 1904, parasites of birds in the Australasian, Nearctic and Neotropical Regions, pp. 1-73 in Zootaxa 5173 (1) on page 51, DOI: 10.11646/zootaxa.5173.1.1, http://zenodo.org/record/696607

FIGURE 15 Ixodes brevisetosus n in Description of eight new species of Ixodes Latreille, 1795 (Acari: Ixodidae) and redescription of I. auritulus Neumann, 1904, parasites of birds in the Australasian, Nearctic and Neotropical Regions

FIGURE 15 Ixodes brevisetosus n. sp., scanning electron micrographs of larva (12 miles east of Edie Creek Road, Bulldog Road, Morobe, Papua New Guinea, USNMENT 01481760). A, Alloscutum; B, Idiosoma, ventral view. Scale-bars: 0.1 mm. SEMs by D.A. ApanaskevichPublished as part of Apanaskevich, Dmitry A., Apanaskevich, Maria A., Klimov, Pavel B., Edgy, Brett M., Bermúdez, Sergio E., Labruna, Marcelo B., Korzeev, Andrei I. & Barker, Stephen C., 2022, Description of eight new species of Ixodes Latreille, 1795 (Acari: Ixodidae) and redescription of I. auritulus Neumann, 1904, parasites of birds in the Australasian, Nearctic and Neotropical Regions, pp. 1-73 in Zootaxa 5173 (1) on page 25, DOI: 10.11646/zootaxa.5173.1.1, http://zenodo.org/record/696607

FIGURE 13. Ixodes brevisetosus n in Description of eight new species of Ixodes Latreille, 1795 (Acari: Ixodidae) and redescription of I. auritulus Neumann, 1904, parasites of birds in the Australasian, Nearctic and Neotropical Regions

FIGURE 13. Ixodes brevisetosus n. sp., scanning electron micrographs of nymph (12 miles east of Edie Creek Road, Bulldog Road, Morobe, Papua New Guinea, USNMENT 01481774). A, Scutum; B, Alloscutum; C, Idiosoma, ventral view; D, Spiracular plate (arrows show orientation of spiracular plate: a, anterior; d, dorsal). Scale-bars: 0.1 mm. SEMs by D.A. ApanaskevichPublished as part of <i>Apanaskevich, Dmitry A., Apanaskevich, Maria A., Klimov, Pavel B., Edgy, Brett M., Bermúdez, Sergio E., Labruna, Marcelo B., Korzeev, Andrei I. & Barker, Stephen C., 2022, Description of eight new species of Ixodes Latreille, 1795 (Acari: Ixodidae) and redescription of I. auritulus Neumann, 1904, parasites of birds in the Australasian, Nearctic and Neotropical Regions, pp. 1-73 in Zootaxa 5173 (1)</i> on page 23, DOI: 10.11646/zootaxa.5173.1.1, <a href="http://zenodo.org/record/6966076">http://zenodo.org/record/6966076</a&gt

FIGURE 16. Ixodes brevisetosus n in Description of eight new species of Ixodes Latreille, 1795 (Acari: Ixodidae) and redescription of I. auritulus Neumann, 1904, parasites of birds in the Australasian, Nearctic and Neotropical Regions

FIGURE 16. Ixodes brevisetosus n. sp., scanning electron micrographs of larva (12 miles east of Edie Creek Road, Bulldog Road, Morobe, Papua New Guinea, USNMENT 01481760). A, Scutum; B, Gnathosoma, dorsal view; C, Gnathosoma, ventral view; D, Gnathosoma, anteroventral view; E, Coxae and trochanters. Scale-bars: A, E, 0.1 mm; B–D, 0.05 mm. SEMs by D.A. ApanaskevichPublished as part of <i>Apanaskevich, Dmitry A., Apanaskevich, Maria A., Klimov, Pavel B., Edgy, Brett M., Bermúdez, Sergio E., Labruna, Marcelo B., Korzeev, Andrei I. & Barker, Stephen C., 2022, Description of eight new species of Ixodes Latreille, 1795 (Acari: Ixodidae) and redescription of I. auritulus Neumann, 1904, parasites of birds in the Australasian, Nearctic and Neotropical Regions, pp. 1-73 in Zootaxa 5173 (1)</i> on page 27, DOI: 10.11646/zootaxa.5173.1.1, <a href="http://zenodo.org/record/6966076">http://zenodo.org/record/6966076</a&gt

FIGURE 36. Ixodes tinamou n in Description of eight new species of Ixodes Latreille, 1795 (Acari: Ixodidae) and redescription of I. auritulus Neumann, 1904, parasites of birds in the Australasian, Nearctic and Neotropical Regions

FIGURE 36. Ixodes tinamou n. sp., scanning electron micrographs of female (Tinki, Ocongate, Cuzco, Peru, USNMENT 00987216, paratype). A, Idiosoma, dorsal view; B, Scutum, dorsal view; C, Scutum, dorsolateral view; D, Alloscutal setae in central area of alloscutum; E, Alloscutal setae in area of alloscutum medial to marginal grooves; F, Idiosoma, ventral view. Scale-bars: A, F, 0.5 mm; B, C, 0.2 mm; D, E, 0.05 mm. SEMs by D.A. ApanaskevichPublished as part of Apanaskevich, Dmitry A., Apanaskevich, Maria A., Klimov, Pavel B., Edgy, Brett M., Bermúdez, Sergio E., Labruna, Marcelo B., Korzeev, Andrei I. & Barker, Stephen C., 2022, Description of eight new species of Ixodes Latreille, 1795 (Acari: Ixodidae) and redescription of I. auritulus Neumann, 1904, parasites of birds in the Australasian, Nearctic and Neotropical Regions, pp. 1-73 in Zootaxa 5173 (1) on page 65, DOI: 10.11646/zootaxa.5173.1.1, http://zenodo.org/record/696607

FIGURE 12. Ixodes brevisetosus n in Description of eight new species of Ixodes Latreille, 1795 (Acari: Ixodidae) and redescription of I. auritulus Neumann, 1904, parasites of birds in the Australasian, Nearctic and Neotropical Regions

FIGURE 12. Ixodes brevisetosus n. sp., scanning electron micrographs of female (12 miles east of Edie Creek Road, Bulldog Road, Morobe, Papua New Guinea, USNMENT 01481776, paratype). A, Spiracular plate (arrows show orientation of spiracular plate: a, anterior; d, dorsal); B, Gnathosoma, dorsal view; C, Gnathosoma, ventral view; D, Gnathosoma, anteroventral view; E, Palpal segment I, medial view; F, Coxae I–III and trochanters I–III; G, Coxa IV and trochanter IV. Scale-bars: A–E, 0.1 mm; F, G, 0.2 mm. SEMs by D.A. ApanaskevichPublished as part of Apanaskevich, Dmitry A., Apanaskevich, Maria A., Klimov, Pavel B., Edgy, Brett M., Bermúdez, Sergio E., Labruna, Marcelo B., Korzeev, Andrei I. & Barker, Stephen C., 2022, Description of eight new species of Ixodes Latreille, 1795 (Acari: Ixodidae) and redescription of I. auritulus Neumann, 1904, parasites of birds in the Australasian, Nearctic and Neotropical Regions, pp. 1-73 in Zootaxa 5173 (1) on page 22, DOI: 10.11646/zootaxa.5173.1.1, http://zenodo.org/record/696607