A new species of soft tick from dry tropical forests of Brazilian Caatinga.

Ornithodoros tabajara n. sp. is described from laboratory-reared larvae and adult specimens collected in the Brazilian Caatinga. This new species shares the ecological niche with Ornithodoros rietcorreai and is likely associated with colonial rodents of genus Kerodon. However, O. tabajara n. sp. is morphologically easy to distinguish from O. rietcorreai and other Neotropical Ornithodoros by a unique combination of characters: larva with 17 pairs of dorsal setae (seven anterolateral, three central and seven posterolateral), sub-oval dorsal plate, hypostome blunt apically with dentition formula 2/2 along its extension, only one pair of posthypostomal setae, six pairs of sternal setae, posteromedian setae absent, and leave-shaped anal valves; alive adults with whitish islands of mammillae symmetrically distributed on dorsum (not visible in ethanol-preserved specimens), and median disk merging with posteromedian file. A phylogenetic analysis performed with mitochondrial 16S rDNA sequences points O. tabajara n. sp. as O. rietcorreai's sister taxon, which rises the hypothesis of sympatric speciation.

Nuclear (18S-28S rRNA) and mitochondrial genome markers of Carios (Carios) vespertilionis (Argasidae) support Carios Latreille, 1796 as a lineage embedded in the Ornithodorinae: re-classification of the Carios sensu Klompen and Oliver (1993) clade into its respective subgenera.

Argasid systematics remains controversial with widespread adherence to the Hoogstraal (1985) classification scheme, even though it does not reflect evolutionary relationships and results in paraphyly for the main genera of soft ticks (Argasidae), namely Argas and Ornithodoros. The alternative classification scheme, proposed by Klompen and Oliver (1993), has problems of its own: most notably paraphyly of the subgenus Pavlovskyella and the controversial grouping together of the subgenera Alectorobius, Antricola, Carios, Chiropterargas, Nothoaspis, Parantricola, Reticulinasus and Subparmatus into the genus Carios. Recent phylogenetic analyses of 18S/28S rRNA sequences and mitochondrial genomes agree with the scheme of Klompen and Oliver (1993), with regard to the paraphyly of Pavlovskyella, placement of Alveonasus, Ogadenus, Proknekalia and Secretargas in the Argasinae and placement of Carios and Chiropterargas in the Ornithodorinae (Mans et al., 2019). The Carios clade and its constituent subgenera remain controversial, since the phylogenetic position of its type species Carios (Carios) vespertilionis Latreille, 1796 (formerly Argas vespertilionis) has not been determined with confidence. The current study aimed to resolve Carios sensu lato Klompen and Oliver, 1993, and Carios sensu stricto Hoogstraal, 1985, by determining and analysing phylogenetic nuclear and mitochondrial markers for C. (C.) vespertilionis. Both the nuclear and mitochondrial markers support placement of Carios s.s. within the subfamily Ornithodorinae, but to the exclusion of the clade that includes the 6 other subgenera that are part of Carios s.l. Klompen and Oliver (1993), namely Alectorobius, Antricola, Nothoaspis, Parantricola, Reticulinasus and Subparmatus. These 6 subgenera form a monophyletic clade that might be placed as new subgenera within the genus Alectorobius, or elevated to genera. Given the substantial differences in biology among these subgenera, we propose that these 6 subgenera be elevated to genera. Thus, we propose to modify the classification scheme of Mans et al. (2019) so that the subfamily Argasinae now has six genera, Alveonasus, Argas (subgenera Argas and Persicargas), Navis, Ogadenus, Proknekalia and Secretargas, and the subfamily Ornithodorinae has nine genera, Alectorobius, Antricola (subgenera Antricola and Parantricola), Carios, Chiropterargas, Nothoaspis, Ornithodoros (subgenera Microargas, Ornamentum, Ornithodoros, Pavlovskyella and Theriodoros), Otobius, Reticulinasus and Subparmatus (genera indicated in bold).

Ticks (Acari: Ixodoidea) associated with mammals in Colombia: a historical review, molecular species confirmation, and establishment of new relationships.

Ticks are considered the second most important vectors of pathogens worldwide, after mosquitoes. This study provides a systematic review of vector-host relationships between ticks and mammals (domestic and wild) and consolidates information from studies conducted in Colombia between 1911 and 2020. Using the PRISMA method, 71 scientific articles containing records for 51 tick species (Argasidae and Ixodidae) associated with mammals are reported. The existing information on tick-mammal associations in Colombia is scarce, fragmented, or very old. Moreover, 213 specimens were assessed based on morphological and molecular analyses, which allowed confirming eight tick species associated with mammals: Amblyomma calcaratum, Amblyomma dissimile, Amblyomma mixtum, Amblyomma nodosum, Amblyomma ovale, Amblyomma varium, Ixodes luciae, and Ixodes tropicalis. Several tick species are molecularly confirmed for Colombia and nine new relationships between ticks and mammals are reported. This research compiles and confirms important records of tick-mammal associations in Colombia.

First Record of Carios kelleyi (Acari: Ixodida: Argasidae) in New Jersey, United States and Implications for Public Health.

The soft tick Carios kelleyi (Cooley and Kohls), a parasite of bats known to occur in at least 29 of the 48 conterminous U.S. states, is here reported from New Jersey for the first time, based on larvae collected from big brown bats, Eptesicus fuscus. Although thought to be widespread in North America, the ecology of C. kelleyi is not well understood, despite reports of this species feeding on humans and its consequent potential as a disease vector. The association of C. kelleyi with bat species that regularly roost in human-made structures, such as attics and barns, and recent isolations from this tick of pathogens capable of infecting humans, companion animals, and livestock underscore the need for further studies of these bat ectoparasites.

Phylogenies from mitochondrial genomes of 120 species of ticks: Insights into the evolution of the families of ticks and of the genus Amblyomma.

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.

Retrospective and new records of ticks (Acari: Argasidae, Ixodidae) from the state of Maranhão, an Amazon-Cerrado transition area of Brazil.

The Brazilian state of Maranhão is located in a transition area of the Amazon and Cerrado biomes, where there is a rich fauna of vertebrates. This study aimed to update the list of the ticks occurring in Maranhão, through a compilation of literature records and examination of three tick collections, plus the addition of unpublished collections of ticks from road-killed animals during recent years. Our results indicate that the tick fauna of Maranhão includes 26 species: seven in the family Argasidae (Antricola guglielmonei, Argas miniatus, Ornithodoros cavernicolous, Ornithodoros hasei, Ornithodoros mimon, Ornithodoros rietcorreai and Ornithodoros rudis); and 19 in the family Ixodidae (Amblyomma auricularium-provisional, Amblyomma cajennense sensu stricto, Amblyomma calcaratum, Amblyomma dissimile, Amblyomma geayi, Amblyomma longirostre, Amblyomma naponense, Amblyomma nodosum, Amblyomma oblongoguttatum, Amblyomma ovale, Amblyomma parvum, Amblyomma rotundatum, Amblyomma sculptum, Amblyomma triste, Dermacentor nitens, Ixodes luciae, Haemaphysalis juxtakochi, Rhipicephalus microplus, and Rhipicephalus sanguineus sensu lato). Eleven of the above species are reported for the first time in Maranhão. We consider previous reports of Ornithodoros talaje and Amblyomma pseudoconcolor in Maranhão as misidentification with O. hasei and A. auricularium, respectively. Until 1958, only 11 tick species were reported in Maranhão, with no additional reports until 2009. During 2010-2019, 15 additional species were reported. We also highlight the importance of ticks for public and veterinary health in Maranhão.

A third species of Nothoaspis Keirans & Clifford, 1975 (Acari: Argasidae): Nothoaspis setosus (Kohls, Clifford & Jones, 1969) n. comb.

In South America, early descriptions of soft tick species were based on examination of the external morphology of the larval stages. In many cases, specimens were collected only once as it is the case of some bat-associated Ornithodoros spp. If we are to understand the systematic scenario of South American soft ticks, these species become axial questions to be re-studied from a morphological and molecular point of view. The objective of this study was to assess the taxonomic identity of soft tick larvae collected on bats inhabiting crevices of a large rock in the Rondônia State (RO), Brazilian Amazon. After a detailed morphological analysis using light microscopy, three large engorged larvae sharing the same phenotype were identified as Ornithodoros setosus Kohls, Clifford & Jones, 1969, a species collected in 1964 on bats in RO. Remarkably, maximum parsimony and Bayesian phylogenetic analyses based on tick 16S rRNA mitochondrial gene sequences obtained from two of these specimens showed that O. setosus indeed corresponds to a species of Nothoaspis Keirans & Clifford, 1975. Therefore, a new combination, Nothoaspis setosus (Kohls, Clifford & Jones, 1969), is herein proposed. While an elongated triangular dorsal plate with a curvy-notched posterior margin, and bulges in the lateral margins of basis capitulum correspond to common characters in larvae of the genus Nothoaspis, polymorphic traits are represented by minute cornua in the basis of the capitulum, the dentition of the hypostomal tip, triangular spurs on coxae I, and the number of dorsal and circumanal setae.

Argasid and ixodid systematics: Implications for soft tick evolution and systematics, with a new argasid species list.

The systematics of the genera and subgenera within the soft tick family Argasidae is not adequately resolved. Different classification schemes, reflecting diverse schools of scientific thought that elevated or downgraded groups to genera or subgenera, have been proposed. In the most recent classification scheme, Argas and Ornithodoros are paraphyletic and the placement of various subgenera remains uncertain because molecular data are lacking. Thus, reclassification of the Argasidae is required. This will enable an understanding of soft tick systematics within an evolutionary context. This study addressed that knowledge gap using mitochondrial genome and nuclear (18S and 28S ribosomal RNA) sequence data for representatives of the subgenera Alectorobius, Argas, Chiropterargas, Ogadenus, Ornamentum, Ornithodoros, Navis (subgen. nov.), Pavlovskyella, Persicargas, Proknekalia, Reticulinasus and Secretargas, from the Afrotropical, Nearctic and Palearctic regions. Hard tick species (Ixodidae) and a new representative of Nuttalliella namaqua (Nuttalliellidae), were also sequenced with a total of 83 whole mitochondrial genomes, 18S rRNA and 28S rRNA genes generated. The study confirmed the utility of next-generation sequencing to retrieve systematic markers. Paraphyly of Argas and Ornithodoros was resolved by systematic analysis and a new species list is proposed. This corresponds broadly with the morphological cladistic analysis of Klompen and Oliver (1993). Estimation of divergence times using molecular dating allowed dissection of phylogeographic patterns for argasid evolution. The discovery of cryptic species in the subgenera Chiropterargas, Ogadenus and Ornithodoros, suggests that cryptic speciation is common within the Argasidae. Cryptic speciation has implications for past biological studies of soft ticks. These are discussed in particular for the Ornithodoros (Ornithodoros) moubata and Ornithodoros (Ornithodoros) savignyi groups.

A checklist of the ticks (Acari: Argasidae, Ixodidae) of Japan.

Ticks significantly affect the health of humans and animals in Japan. However, no complete checklist has been produced for the Japanese tick fauna in English, and even recent checklists in Japanese do not include all presently known tick species found on the Japanese islands. Here, a complete species list of all tick taxa known to occur within Japan is presented. The status of several controversial Japanese tick species is also discussed.

The fossil record and the origin of ticks revisited.

The fossil record of ticks has been classically used to discern the probable vicariance events of these arthropods, and to date the split moments of the different lineages. Although new techniques based on molecular clock methods are already available, tick fossil record still stands as a valuable source of information if correctly interpreted. Here we report about a male specimen of Ornithodoros (Alectorobius) sp. found in Dominican amber of about 25 millions years, illustrating its main morphological features. We also discuss the findings in a recent paper, which may contain unreliable interpretations on some fossil ticks. We support previous data on the presence of Argasidae in Miocene Dominican amber, and provide comments on the presence of ticks in Burmese amber. We further elaborate on the spread of ticks in the light of the record and description of a new family found in Cretaceous amber.

The geographic distribution of Argas (Persicargas) miniatus and Argas (Persicargas) persicus (Acari: Argasidae) in America, with morphological and molecular diagnoses from Brazil, Chile and Cuba.

High similarity of morphological traits has historically overshadowed the identities and distributions of poultry-associated soft ticks Argas (Persicargas) miniatus and Argas (Persicargas) persicus in America. In order to model the occurrence of both parasites in the continent, in the current study we performed morphological and molecular analyses to identify ticks collected in hen houses from Brazil and northern Chile. Combining these results with literature data, and the examination of Argas allotments deposited in the tick collections "Coleção Nacional de Carrapatos Danilo Gonçalves Saraiva" (Brazil), the "Coleção Acarológica do Instituto Butantan São Paulo" (Brazil), and the "Colección Zoológica de la Academia de Ciencia de Cuba" (Cuba), we present a critical list with the localities where A. (P.) miniatus and A. (P.) persicus have been reported in the American continent. Our results confirmed the presence of A. (P.) miniatus in Brazil and Cuba, and A. (P.) persicus in Chile, which in particular, constitutes the first molecularly confirmed report of the later species for South America. Although A. (P.) miniatus and A. (P.) persicus have been documented in 21 American countries, the identity of some reports must still be considered as uncertain until detailed morphological and/or molecular studies are performed. When contrasted to a Köppen-Geiger climate classification, A. (P.) miniatus predominantly occurs in equatorial and A. (P.) persicus in arid climates. However, until undetermined reports of both species are correctly identified, any conclusion on their geo-climatological occurrence throughout the American continent would be rather speculative.

Guia de vigilância acarológica: vetores e hospedeiros da febre maculosa e outras riquetsioses no Rio Grande do Sul / Acarological surveillance guide for vectors and hosts of spotted fever and other rickettsial in Rio Grande do Sul

Estudo dos carrapatos como vetor de doenças aos humanos.

Ixodoidea of the Western Palaearctic: A review of available literature for identification of species.

We aim to produce a review of the most important literature references necessary for the identification of ticks of the families Ixodidae and Argasidae in Europe and northern Africa (i.e. the Western Palaearctic region). The purpose of this paper is to pinpoint a set of critically selected papers that contain reliable information on morphology, taxonomic keys, and comparative discussions which are critical for the identification of the ticks reported in the target region. When necessary, comments are provided on the systematic position of a species, or on suitable papers already addressing the issue. This review includes a list of 216 references which cover all Ixodoidea species reported as permanent residents of the Western Palaearctic, namely 28 species of the genus Ixodes, two Dermacentor, seven Haemaphysalis, nine Hyalomma, eight Rhipicephalus, five Argas and about seven species of Ornithodoros.

A new species of Ornithodoros (Acari: Argasidae) from desert areas of northern Chile.

Ornithodoros atacamensis n. sp. is described from larvae collected on the lizard Liolaemus bisignatus and from free-living adults collected in desert areas from the Pan de Azúcar and Llanos de Challe National Parks, in Northern Chile. Additionally, unengorged larvae were obtained from fertilized females, which laid eggs in the laboratory. Morphological and mitochondrial 16S rDNA sequence analyses were performed in order to compare this new soft tick species with other congeneric Neotropical representatives. Larvae of O. atacamensis are morphologically closely related to Ornithodoros talaje sensu stricto, Ornithodoros puertoricensis, Ornithodoros rioplatensis, Ornithodoros guaporensis and Ornithodoros hasei, all belonging to the O. talaje species group. The larval diagnostic characters for this species are a combination of a large pyriform dorsal plate with a length of approximately 300µm, 17 pairs of dorsal setae with five central pairs, hypostome with apex pointed and dental formula 2/2 in most rows, 3/3 apically, and capsule of the Haller's organ oval in shape without reticulations. Phylogenetic analyses inferred from the mitochondrial 16S rRNA gene and a Principal Component Analysis based on morphometric characters provide additional support to the description of O. atacamensis as an independent lineage within the genus clustering within the O. talaje species group.

Redescription of Ornithodoros dyeri (Ixodida: Argasidae) based on morphologic and molecular data.

Larvae, nymphs and adults of the cave dwelling tick Ornithodoros dyeri, collected in 3 Mexican states, were studied using morphological and molecular methods. The adults and nymphs were characterized by an elongated body in proportion to the width and a dorsum bounded by two contiguous ridges and one third ridge (inner) that was incomplete on each side. The larvae of this species have 14 pairs of dorsal setae, a venter body with nine pairs of setae plus a posteromedian; a moderately large, dorsal plate and piriform, a hypostome arising from a relatively short, subtriangular median extension of the basis capituli, and a capsule of Hallers organ with reticulations. Based on a maximum likelihood analysis of the sequences of a fragment of approximately 414bp of the mitochondrial 16S rRNA gene, we showed that O. dyeri represents an independent lineage within neotropical species of the Argasidae. The bat species Mimon cozumelae and Peropteryx macrotis represents a new host record for this argasid.

An overview on Sardinia's soft ticks (Ixodida: Argasidae).

Knowledge about soft ticks (Ixodida: Argasidae) in Sardinia is incomplete and distribution data need to be updated. This work studies soft ticks on the island focusing on two species, Argas reflexus and Ornithodoros maritimus, both recently recorded. A total number of 12 specimens of these species of interest were collected between 2004 and 2015. This study reports for the first time the presence of O. maritimus in a coastal area in Italy, and more generally in a coastal area rather than small islands near the coastline, confirming the presence of this species on the island 20 years after its last recording. Moreover we confirm the presence of A. reflexus on the island, in the town of Cagliari and, for the first time, in the town of Quartu Sant'Elena. At the present state of knowledge, in Sardinia, Ornithodoros erraticus, which was actively looked for within the surveillance for African swine fever, an endemic disease since 1978 on the island, is not present. The presence of another species reported only once in Sardinia, Argas vespertilionis, needs further confirmation.

Detection of a novel Rickettsia sp. in soft ticks (Acari: Argasidae) in Algeria.

Argasid ticks are vectors of viral and bacterial agents that can infect humans and animals. In Africa, relapsing fever borreliae are neglected arthropod-borne pathogens that cause mild to deadly septicemia and miscarriage. It would be incredibly beneficial to be able to simultaneous detect and identify other pathogens transmitted by Argasid ticks. From 2012 to 2014, we conducted field surveys in 4 distinct areas of Algeria. We investigated the occurrence of soft ticks in rodent burrows and yellow-legged gull (Larus michahellis) nests in 10 study sites and collected 154 soft ticks. Molecular identification revealed the occurrence of two different soft tick genera and five species, including Carios capensis in yellow-legged gull nests and Ornithodoros occidentalis, Ornithodoros rupestris, Ornithodoros sonrai, Ornithodoros erraticus in rodent burrows. Rickettsial DNA was detected in 41/154, corresponding to a global detection rate of 26.6%. Sequences of the citrate synthase (gltA) gene suggest that this agent is a novel spotted fever group Rickettsia. For the first time in Algeria, we characterize a novel Rickettsia species by molecular means in soft ticks.

The ticks (Acari: Ixodida: Argasidae, Ixodidae) of Bolivia.

The tick species reported in Bolivia are reviewed here as (1) endemic or established: Ornithodoros echimys, O. guaporensis, O. hasei, O. kohlsi, O. mimon, O. peropteryx, O. rostratus, Otobius megnini, Amblyomma auricularium, A. cajennense, A. calcaratum, A. coelebs, A. dubitatum, A. humerale, A. incisum, A. longirostre, A. naponense, A. nodosum, A. oblongoguttatum, A. ovale, A. parvitarsum, A. parvum, A. pecarium, A. pseudoconcolor, A. rotundatum, A. scalpturatum, A. tigrinum, A. triste, Dermacentor nitens, Haemaphysalis juxtakochi, H. leporispalustris, I. boliviensis, I. cooleyi, I. luciae, Rhipicephalus microplus, R. sanguineus, and (2) erroneously reported: Ornithodoros puertoricensis, O. talaje, O. turicata, Amblyomma americanum, A. maculatum, A. multipunctum, Ixodes ricinus, I. scapularis, Rhipicephalus annulatus. Many of these records are lacking locality and/or host, and some of them need new findings for confirmation. Some of the species recorded may represent a threat for human and animal health, therefore would be of great value to make a countrywide survey of ticks in order to update the information presented in this work.

Immature argasid ticks: diagnosis and keys for neotropical region / Carrapatos argasídeos imaturos: diagnoses e chaves para a região neotropical

Many argasid tick species are known only through their larval descriptions, in which the chaetotaxy, together with other external morphological characteristics, has been used to separate genera and species. However, the illustrations of these features are based on optical microscopy alone and many of these features are not clearly defined. Because of the difficulties in determining the larval and nymph stages of some genera, we have prepared illustrated keys for the immature stages of argasids, including an up-to-date list of the known species of the Neotropical region. We have also included an illustrated key for larvae of the Ornithodoros species from Brazil, based on scanning electron microscopy.

Muitos carrapatos argasídeos são conhecidos somente por descrições larvais, nas quais a quetotaxia associada a outros caracteres morfológicos tem sido usada para separar gêneros e espécies. No entanto, as ilustrações sobre esses caracteres são baseadas somente em microscopia óptica e muitos deles não estão claramente definidos. Devido às dificuldades em determinar estágios larvais e ninfais de alguns gêneros, elaboramos chaves ilustradas para os estágios imaturos de argasídeos, incluindo uma lista atualizada de espécies conhecidas da região Neotropical. Incluímos também uma chave ilustrada para larvas das espécies de Ornithodoros do Brasil baseada em microscopia eletrônica de varredura.