RESUMO
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.
RESUMO
The evolution and phylogenetic relationships of the ticks at both the family and genus levels are contested. The genus Amblyomma and its subgenera are in a state of flux; moreover, the relationships among the three tick families are controversial due to conflicting phylogenetic support for different arrangements of the three families of living ticks. With 18 newly sequenced mitochondrial (mt) genomes of ticks included, we executed the largest mt genome phylogenetic study of ticks so far. Phylogenetic trees were inferred from one sea spider mt genome, one horseshoe crab, five mite mt genomes and 146 tick mt genomes from 120 species: 153 mt genomes in total. Sixteen phylogenetic trees were inferred from 10 datasets using both maximum likelihood and Bayesian inference methods. We describe the first novel mt gene-arrangement for the metastriate Ixodidae in Amblyomma (Africaniella) transversale. Also, three unusual partial 16S rRNA gene inserts were found in the mt genome of Haemaphysalis (Alloceraea) kitaokai: we consider the possible role of past genome translocation events in the formation of these inserts. Our phylogenies revealed evidence that: (i) the genus Amblyomma is polyphyletic with respect to Amblyomma (Africaniella) transversale; (ii) the subgenus Aponomma is apparently embedded in the genus Amblyomma; (iii) Haemaphysalis (Segalia) parva and Haemaphysalis (Alloceraea) kitaokai form a clade to the exclusion of other Haemaphysalis species; and (iv) the phylogenetic position of the family Nuttalliellidae is unstable among phylogenies from different datasets.
Assuntos
Amblyomma/genética , Argasidae/classificação , Evolução Biológica , Genoma Mitocondrial , Ixodidae/classificação , Filogenia , AnimaisRESUMO
Seventy species of ticks are known from Australia: 14 soft ticks (family Argasidae) and 56 hard ticks (family Ixodidae). Sixteen of the 70 ticks in Australia may feed on humans and domestic animals (Barker and Walker 2014). The other 54 species of ticks in Australia feed only on wild mammals, reptiles and birds. At least 12 of the species of ticks in Australian also occur in Papua New Guinea. We use an image-matching system much like the image-matching systems of field guides to birds and flowers to identify Ixodes holocyclus (paralysis tick), Ixodes cornuatus (southern paralysis tick) and Rhipicephalus (Boophilus) australis (Australian cattle tick). Our species accounts have reviews of the literature on I. holocyclus (paralysis tick) from the first paper on the biology of an Australian tick by Bancroft (1884), on paralysis of dogs by I. holocyclus, to papers published recently, and of I. cornuatus (southern paralysis tick) and Rhipicephalus (Boophilus) australis (Australian cattle tick). We comment on four controversial questions in the evolutionary biology of ticks: (i) were labyrinthodont amphibians in Australia in the Devonian the first hosts of soft, hard and nuttalliellid ticks?; (ii) are the nuttalliellid ticks the sister-group to the hard ticks or the soft ticks?; (iii) is Nuttalliella namaqua the missing link between the soft and hard ticks?; and (iv) the evidence for a lineage of large bodied parasitiform mites (ticks plus the holothyrid mites plus the opiliocarid mites).