Ticks (Acari: Argasidae and Ixodidae) infesting

Systematic & Applied Acarology 23(8): 1497–1508 (2018) http://doi.org/10.11158/saa.23.8.1 Article

ISSN 1362-1971 (print) ISSN 2056-6069 (online)

http://zoobank.org/urn:lsid:zoobank.org:pub:440D267D-54DD-4A62-A14F-66587A98F7FD

Ticks (Acari: Argasidae and Ixodidae) infesting amphibians and reptiles in Northeastern Brazil EDNA PAULINO DE ALCANTARA1, CRISTIANA FERREIRA-SILVA1, ROBSON WALDEMAR ÁVILA2, RICHARD DE CAMPOS PACHECO3, THIAGO FERNANDES MARTINS4, SEBASTIÁN MUÑOZ-LEAL4 & DRAUSIO HONORIO MORAIS5 1

Programa de Pós-Graduação em Ciências Biológicas (Zoologia), Departamento de Parasitologia, Laboratório de Parasitologia de Animais Silvestres, Instituto de Biociências, Universidade Estadual Paulista – UNESP, Av. Bento Lopes s/n Distrito de Rubião Junior CEP 18080-970, Botucatu, São Paulo, Brazil. 2 Programa de Pós-Graduação em Bioprospecção Molecular, Universidade Regional do Cariri, Departamento de Química Biológica, Laboratório de Herpetologia, Rua Cel. Antônio Luiz, 1161, CEP 63100-000, Crato, Ceará, Brazil. 3 Universidade Federal de Mato Grosso, Faculdade de Medicina Veterinária – FAVET, Programa de Pós-graduação em Ciências Veterinárias-PPGVET, Av. Fernando Corrêa da Costa, 2.367, Boa Esperança, CEP 78060-900, Cuiabá, Mato Grosso, Brazil. 4 Departamento de Medicina Veterinária Preventiva e Saúde Animal, Faculdade de Medicina Veterinária e Zootecnia, Universidade de São Paulo, Av. Prof. Orlando Marques de Paiva, 87, Cidade Universitária, CEP 05508-270, São Paulo, São Paulo, Brazil. 5 Universidade Federal Rural da Amazônia – UFRA, PA 275, Km 13, Zona Rural. CEP 68515-000, Parauapebas, Pará, Brasil. Corresponding author: Edna Paulino de Alcantara, Tel: +55 14 981371866 E-mail: [email protected]

Abstract Ticks collected on 21 different species of amphibians and reptiles captured during 2001 to 2017 in the northeast Brazil were identified. Overall, 227 individuals (185 larvae, 32 nymphs, 7 adults females and 3 males) identified as follow: Amblyomma dissimile, Amblyomma rotundatum, Amblyomma sp. larvae, Ornithodoros mimon, Ornithodoros rietcorreai, and an Ornithodoros sp. Herein, we present 25 new tick-host associations, thus contributing to the knowledge on tick-fauna associated to amphibians and reptiles from Brazilian northeast. Keywords: Amblyomma, Ornithodoros, ectoparasites

Introduction Ticks are a worldwide-distributed group of invertebrates that parasitizes a broad variety of hosts, ranging from small amphibians, reptiles and birds to large mammals, including humans (Jongejan & Uilenberg 2004, Guglielmone et al. 2006, Nava et al. 2007, Ogrzewalska & Pinter 2016). Recent taxonomical studies on ticks from the Neotropical region have suggested that specific richness of these ectoparasites could be underestiamted (Estrada-Peña et al. 2003, Labruna et al. 2008, Venzal et al. 2008, 2012, 2013, Labruna & Venzal 2009, Nava et al. 2010, 2013, 2014a, Dantas-Torres et al. 2012, Krawczak et al. 2015, Venzal et al. 2015, Muñoz-Leal et al. 2016, 2017, Labruna et al. 2016). In Brazil, 72 species of ticks occur distributed in two families: Ixodidae (46 species) and Argasidae (26 species) (Guglielmone & Nava 2010, Dantas-Torres et al. 2012, BarrosBattesti et al. 2013, 2015, Martins et al. 2014, Nava et al. 2014a, Krawczak et al. 2015, Labruna et al. 2016, Wolf et al. 2016, Muñoz-Leal et al. 2017).

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The genus Amblyomma is worldwide represented by 138 valid species (Guglielmone et al. 2014, Nava et al. 2014a, 2014b, Krawczak et al. 2015), from which 64 species are present in the Neotropical zoogeographic region, with 32 recorded in Brazil (Dantas-Torres et al. 2009, Martins et al. 2010, Nava et al. 2014a, Krawczak et al. 2015). Four species have been recorded in Brazilian amphibians and reptiles (Amblyomma auricularium (Conil 1878), Amblyomma fuscum Neumann, 1907, Amblyomma dissimile Koch, 1844, Amblyomma rotundatum Koch, 1844, Amblyomma humerale Koch, 1844, Amblyomma parvum Aragão, 1908, and Amblyomma sculptum Berlese, 1888 (Sinkoc & Brum 1997, Barros-Battesti et al. 2006, Martins et al. 2007, Dantas-Torres et al. 2008, 2010, Morais et al. 2010, 2017, Martins et al. 2015,Kluyber et al. 2016, Oda et al. 2018) with three records in the northeastern region of the country (Dantas-Torres et al. 2008, 2010, Quirino et al. 2016). The genus Ornithodoros is currently represented by 134 species, with 61 representatives in the Neotropical zoogeographic region, and 21 in Brazil (Dantas-Torres et al. 2012, Heath 2012, Venzal et al. 2013, Barros-Battesti et al. 2015, Wolf et al. 2016, Muñoz-Leal et al. 2017, Bakkes et al. 2018). Recently, two species of Ornithodoros associated with amphibians inhabiting in remnants of the Atlantic Rainforest biome have been described (Barros-Battesti et al. 2015, Muñoz-Leal et al. 2017). However, records of argasid ticks parasitizing reptiles across Brazilian biomes are scarce, and reduced to a sole study in which larval and post-larval stages of an unidentified Ornithodoros sp. were reported in association with a captive Boa constrictor constrictor Linnaeus, 1758 in Rio Grande do Norte state, Brazilian northeast (Pereira et al. 2012). According to Dantas-Torres et al. (2008), there is limited information on taxonomy and ecology of ticks infesting ectothermic animals in the Northeastern Brazil. In order to contribute with the knowledge of ticks infesting Brazilian herpetozoans, in the present study we analyzed museum specimens collected in the Brazilian northeast in search for ticks and described new host-parasite associations. Material and Methods Examined material This study analyzed 131 species (5.312 specimens) of amphibians and 151 species (4.938 specimens) of reptiles deposited in the Coleção Herpetológica da Universidade Regional do Cariri (URCA). The amphibian and reptile species were captured between 2001 and 2017 in 16 municipalities from Alagoas (AL), Ceará (CE), Maranhão (MA), Pernambuco (PE), Piauí (PI) states (Fig. 1). Ticks were collected manually with tweezers and preserved in 70% ethanol for taxonomic identification. Ticks were examined under the stereomicroscope and ixodid species were identified following Aragão and Fonseca (1961), Onofrio et al. (2006), Barros-Battesti et al. (2006) and Martins et al. (2010). Soft ticks were clarified in a 20% KOH solution, mounted in semi-permanent slides using Hoyer’s medium and observed in an Olympus BX40 optical microscope. Subsequently, morphological identification of slide-mounted specimens was performed using the key of Jones and Clifford (1972), and original species descriptions of other Neotropical argasids (Endris et al. 1989, Venzal et al. 2008, Nava et al. 2013, Barros-Battesti et al. 2015, Labruna et al. 2016, Muñoz-Leal et al. 2016, 2017). Results A total of 45 specimens from at least 21 species were sampled: three amphibians and 18 reptiles were parasitized by at least one tick species (Table 1). 1498

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FIGURE 1. Map showing the localities where ticks were sampled in the Northeastern Brazil.

A total of 227 tick specimens (seven females, three males, 32 nymphs, and 185 larvae) belonging to the following six taxa were collected: Amblyomma dissimile (N=6), Amblyomma rotundatum (N=36), Amblyomma sp. larvae (N=92), Ornithodoros mimon Kohls 1969 (N=87), Ornithodoros rietcorreai Labruna, Nava & Venzal 2016 (N=3) and Ornithodoros sp. larvae (N=3) (Table 1). Ethanol preserved specimens were deposited at the collection of Universidade Federal de Mato Grosso (UFMT). Slide-mounted larvae were deposited at the “Coleção Nacional de Carrapatos Danilo Gonçalves Saraiva” of Faculdade de Medicina Veterinária e Zootecnia da Universidade de São Paulo (CNC) under the following accession numbers: CNC-3511, 3512, and 3513. A total of 10 adults, 32 nymphs, and 92 Amblyomma sp. larvae were collected on one species of amphibian, three species of lizards and eight species of snakes (Table 1). A total of 93 Ornithodoros larvae were isolated from four species of snakes (Table 1). Ornithodoros mimon (N=87) was morphologically identified by the following combination of characters: dorsal plate pear-like in shape, 11 pairs of dorsolateral and three pairs of central setae; hypostome uniform in width and apically blunt, with 3–4 row of denticles (19 denticles in row I; 16 in row II, 6–7 in row III, 4–5 in row IV), dentition formula 4/4–3/3 in the apex, and 2/2 towards the base (Kohls et al. 1969). Identification of Ornithodoros rietcorreai (N=3) relied in the presence of the following characters: dorsal surface provided with 11 pairs of dorsolateral and three pairs of central setae; dorsal plate pyriform with a slightly concave posterior margin, anal valves long with leaf-shaped ends (Labruna et al. 2016). Finally, three larvae were identified only as Ornithodoros sp. by sharing the following characters: dorsal plate pyriform, 21 dorsolateral pairs (seven anterolateral, 10 posterolateral) and four central pairs of setae; hypostome long and apically pointed, with three rows of denticles (24 denticles in row I; 21 in row II, and nine in row III); dentition formula 2/2 in the apex, then 3/3 in the middle, and 2/2 towards the base.

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TABLE 1. Ticks collected from amphibians and reptiles in northeastern Brazil. Ticks/Taxon

F

M N

Amblyomma dissimile

2

3

Amblyomma rotundatum

2

L

Host Species

Locality

Ixodidae

1

1

Iguana iguana

Ribeiro Gonçalves—PI

1

Rhinella jimi

Barro, Ipú—CE

1

Rhinella gildae *

São Pedro da Água Branca—MA

2

Rhinella marina

São Pedro da Água Branca—MA

2

Chironius flavolineatus *

Caririaçu, Barbalha—CE

2

Drymarchon corais *

Guaramiranga—CE

1

Lygophis dilepis *

Crato—CE

5

Oxyrhopus trigeminus *

ESEC Aiuaba, Barro—CE

7

Philodryas olfersii

Barro—CE

1

Philodryas nattereri

Barro—CE

1

Xenopholis scalaris *

RPPN - Pedra D'antas, Lagoa dos Gatos—PE

5

Micrurus ibiboboca

ESEC Aiuaba, Barbalha—CE

1

Bothrops leucurus

Crato, Barbalha—CE

2

Bothrops erythromelas *

Farias Brito—CE

1

Copeoglossum arajara *

Ipú—CE

7

Norops brasiliensis *

Crato—CE

1

Chironius exoletus *

Timon—MA

2

Chironius flavolineatus *

Barbalha—CE

1

Erythrolamprus viridis *

Farias Brito—CE

1

Leptodeira annulata *

Varzea Alegre—CE

1

Lygophis dilepis *

Exu—PE

12


Barbalha, Farias Brito—CE; Quebrangulo—AL; Exu—PE

2

Philodryas nattereri *

Exu—PE; Barro—CE

52

Philodryas olfersii

Barro—CE

5

Pseudoboa nigra *

Lavras da Mangabeira—CE; Ouricuri—PE

3

Xenopholis scalaris *

RPPN - Pedra D'antas, Lagoa dos Gatos—PE

2

Micrurus ibiboboca *

Trilha Ecológica do Grangeiro, Crato—CE

2

Bothrops leucurus *

Araripe, Crato—CE

Ornithodoros mimon

87

Corallus hortulanus *

Chapada do Araripe, Crato—CE

Ornithodoros sp.

1


Jati—CE

1


Barro—CE

1

Philodryas olfersii *

Barro—CE

3


Farias Brito—CE

1

1

Amblyomma sp.

Argasidae

Ornithodoros rietcorreai

* New tick-host association, F = Females, M = Males, N = Nymphs, L = Larvae, AL = Alagoas, CE = Ceará, MA = Maranhão, PE = Pernambuco, PI = Piauí.

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Discussion Amblyomma dissimile is an ectoparasite commonly found on amphibians and reptiles, and it has a widespread distribution in the Neotropical region (Fairchild et al. 1966, Jones et al. 1972, Guglielmone et al. 2003). In Brazil record of A. dissimile have been recorded for wild hosts: Rhinella bergi (Céspedez, 2000 “1999”), Rhinella marina (Linnaeus, 1758), Rhinella schneideri (Werner, 1894), Acanthochelys macrocephala (Rhodin, Mittermeier and McMorris, 1984), Chelonoidis carbonarius (Spix, 1824), Chelonoidis denticulatus (Linnaeus, 1766), Kinosternon scorpioides (Linnaeus, 1766), Phrynops geoffroanus (Schweigger, 1812), Rhinoclemmys punctularia (Daudin, 1801), Ameiva ameiva (Linnaeus, 1758), Iguana iguana (Linnaeus, 1758) Tropidurus hispidus (Spix, 1825), Boa constrictor (Stull, 1932), Crotalus durissus cascavella Wagler in Spix, 1824, Chironius laurenti Dixon, Wiest and Cei, 1993, Eunectes murinus (Linnaeus, 1758), Hydrodynastes gigas (Duméril, Bibron & Duméril, 1854), Mastigodryas bifossatus (Raddi, 1820), Mussurana bicolor (Peracca, 1904), Xenodon severus (Linnaeus, 1758) (Schumaker & Barros 1994, Guerra et al. 2000, Brum & Rickes 2003, Freitas et al. 2004, Dantas-Torres et al. 2008, Fischer et al. 2009, Lopes et al. 2010, Scofield et al. 2011, Witter et al. 2016, Nascimento et al. 2017). Amblyomma dissimile has been reported in captive animals Caiman yacare (Daudin, 1801), Iguana iguana, Eunectes notaeus Cope, 1862, Trachemys scripta (Schoepff, 1792), Python reticulatus Schneider, 1801 (exotic snake) (Teixeira et al. 2003, Witter et al. 2016). Amblyomma rotundatum occurs from Florida to Argentina, mainly in ectothermic animals (Oliver et al. 1993, Dantas-Torres et al. 2008). In Brazil, A. rotundatum has been naturally found infesting the species Physalaemus nattereri (Steindachner, 1863), Rhinella arenarum (Hensel, 1867), Rhinella bergi, Rhinella crucifer (Wied-Neuwied, 1821), Rhinella granulosa (Spix, 1824), Rhinella jimi (Stevaux, 2002), Rhinella icterica (Spix, 1824), Rhinella marina, Rhinella sp., Rhinella schneideri, Chelonoidis carbonaria (Spix, 1824), Chelonoidis denticulata (Linnaeus, 1766), Kinosternons scorpioides (Linnaeus, 1766), Mesoclemmys vanderhaegei (Bour, 1973), Platemys platycephala (Schneider, 1792), Ameiva ameiva, Iguana iguana, Tropidurus hispidus, Tropidurus torquatus (Wied, 1820), Tropidurus sp., Plica plica (Linnaeus, 1758), Paleosuchus palpebrosus (Cuvier, 1807), Paleosuchus trigonatus (Schneider, 1801), Boa constrictor, Boa sp., Bothrops alternatus Duméril, Bibron & Duméril, 1854, Bothrops atrox (Linnaeus, 1758), Bothrops jararaca (Wied, 1824) Bothrops jararacussu Lacerda, 1884, Bothrops leucurus Wagler in Spix, 1824, Bothrops moojeni Hoge, 1966, Bothrops neuwiedi Wagler in Spix, 1824, Chironius fuscus (Linnaeus, 1758), Chironius laevicollis (Wied, 1824), Crotalus durissus cascavella, Lachesis muta (Linnaeus, 1766) Micrurus ibiboboca (Merrem, 1820), Philodryas olfersii (Liechtenstein, 1823), Philodryas nattereri Steindachner, 1870, Spilotes pullatus (Linnaeus, 1758) Xenodon neuwiedii Günther, 1863 (Oba & Schumaker 1983, Guerra et al. 2000, Labruna et al. 2002, 2005, Santos et al. 2002, Woehl 2002, Ahid et al. 2003, Brum & Costa 2003, Arzua et al. 2005, Onofrio, 2007, Szabó et al. 2007, Dantas-Torres et al. 2008, 2009, 2010, Pontes et al. 2009, Morais et al. 2010, Rodrigues et al. 2010 Gambale et al. 2014, Guglielmone & Nava 2010, Viana et al. 2012, Luz et al. 2012, 2015, Gomides et al. 2015, Teixeira et al. 2014, Quirino et al. 2016, Witter et al. 2016). Amblyomma rotundatum has been reported in captive animals: Chelonoidis carbonaria, B. constrictor, Corallus hortulanus (Linnaeus, 1758), Crotalus durissus cascavella, Epicrates cenchria (Linnaeus, 1758) Python bivittatus Kuhl 1820 (exotic snake) (Cunha et al. 2003, Dantas-Torres et al. 2005, 2010). Herein, eight new hosts records are provided for nymphal stages of A. rotundatum (Table 1). Thus, severe infestations by A. rotundatum in amphibians and reptiles can cause bleeding and even death (Keirans & Durden 1998). Amblyomma sp. larvae have been found infesting the wild hosts Rhinella schneideri, Ameiva ameiva, Iguana iguana, Tropidurus hispidus, Tropidurus sp., Boa constrictor, Bothrops jararaca, 2018


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Bothrops jararacussu, Chironius fuscus, Chironius laevicollis, Crotalus durissus cascavella, Eunectes murinus, Epicrates cenchria, Philodryas olfersii, Philodryas sp., Spilotes pullatus, Tantilla melanocephala (Linnaeus, 1758), Xenodon neuwiedii (Lizaso 1984, Perreira et al. 2000, Szabó et al. 2007, Dantas-Torres et al. 2008, 2010, Pontes et al. 2009, Lopes et al. 2010, Luz et al. 2012, Witter et al. 2016, Nascimento et al. 2017). Amblyomma sp. larvae have been found infesting the captive animals: Boa constrictor, Epicrates cenchria, Spilotes pullatus (Dantas-Torres et al. 2010). Additionally, in the present study thirteen new host records in reptiles are described for Amblyomma larvae stage (Table 1). In Brazil, there are records of parasitism by argasids of the genus Ornithodoros in the anurans Thoropa miliaris (Spix, 1824) (Barros-Battesti et al. 2015, Sá-Hungaro et al. 2016) and Cycloramphus boraceiensis Heyer, 1983 (Muñoz-Leal et al. 2017), and in the snake B. constrictor constrictor (Pereira et al. 2012). Ornithodoros mimon was originally described from bats in Bolivia and Uruguay (Kohls et al. 1969), yet later records expanded its distribution as a parasite of bats in Argentina and Brazil (Venzal et al. 2004, Barros-Battesti et al. 2006), and also associated to rodents, marsupials, and even humans in Brazil (Labruna et al. 2014, Landulfo et al. 2012). The current record in Corallus hortulanus could be explained by the feeding habits of this snake, which preys mainly on bats, small rodents and marsupials (Santos-Costa et al. 2015). The present study corresponds to the first record of O. mimon parasitizing reptiles, suggesting that this species could also be adapted to feed on ectothermic hosts. Ornithodoros rietcorreai was originally described in Kerodon rupestris Cuvier, 1825 in the Caatinga domain (Labruna et al. 2016). As for O. mimon, the present study records for the first time O. rietcorreai parasitizing reptiles. Moreover, this is also the first record of this species in Ceará State, enlarging its distribution, which is already known for the northeastern Brazilian States of Paraiba and Piauí (Labruna et al. 2016). Although three larvae of Ornithodoros were not identified to species level, their morphological traits were related with specimens of the Ornithodoros talaje group (Guérin-Méneville, 1849). Recent studies have proposed that Ornithodoros talaje is not established in Brazil, noting that previous reports of this species were possibly misidentifications of closely related species yet to be described (Venzal et al. 2008, Labruna et al. 2014). Larvae of Ornithodoros sp. found in the current study could represent a putative new species. However, a detailed morphological study of a larger amount of larvae, the examination of nymphal and adult specimens, and a molecular identification would be necessary in order to confirm this hypothesis. All records of infestation by Ornithodoros mimon, O. rietcorreai and Ornithodoros sp. in snakes from Brazilian northeast constitute new host records (Table 1). Further studies are necessary to know with accuracy the importance of Brazilian snakes as host for Ornithodoros spp. given that such ticks have been previously reported almost exclusively on mammals, including humans. There are still gaps regarding the taxonomy, and host associations of ticks in amphibians and reptiles from Brazil. Therefore, studies on taxonomy, biology and ecology are necessary to quantify more precisely these associations, especially concerning the evolution and transmission of pathogenic agents. The present study expands host records and biogeography of amphibian and reptile associated ticks in the northeast region of Brazil.

Conflict of interest The authors declare no competing personal or financial interests.

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Acknowledgements We thank Fabio Hideki Yamada for manuscript revision. Drausio Honorio Morais thanks Cordenação de Aperfeiçoamento de Pessoal de Nível Superior for providing a research fellowship (CAPES/PNPD 22005013001P4), Robson Waldemar Ávila thanks Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) for providing research fellowship (PQ # 303622/ 2015-6) and Richard de Campos Pacheco thanks CNPq for the scholarship. We are grateful to the CNPq/FUNCAP/CAPES for financial support (PROTAX – Processes 440511/2015-1; 5574685/ 2017; 88882.156872/2016-01; Edital CNPq/ ICMBio Chamada 18/2017. Processo FUNCAP ICM0132-00006.01.00/18).

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simile Koch (Acari: Ixodidae) Atacking Primolius maracana Vieillot (Psitaciformes: Psitacidae) in the Amazon Region, State of Pará, Brazil. Neotropical Entomology, 40, (4), 509–511. http://dx.doi.org/10.1590/S1519-566X2011000400017 Sinkoc, A.L. & Brum, J.G.W. (1997) Ocorrência de Amblyomma fuscum Neumann, 1899 e Amblyomma humerale Koch, 1844 (Acari: Ixodidae) em Bufo arenalis no estado de São Paulo, Brasil. Ciência Rural, 27, (2), 339– 340. https://doi.org/10.1590/S0103-84781997000200029 Szabó, M.P.J., Olegário, M.M.M. & Santos, A.L.Q. (2007) Tick fauna from two locations in the Brazilian savannah. Experimental and Applied Acarology, 43, 73–84. https://doi.org/10.1007/s10493-007-9096-8 Teixeira, G.M.S.L., Gomes, S.L., Dias, M.M.B., Pereira, J.S., Adid, S.M.M. & Freitas, C.I.A. (2014) Detecção de Amblyomma rotundatum (Koch, 1844) em cobra corre campo (Philodryas nattereri, steindachner, 1870). 40º CONBRAVET Congresso Brasileiro de Medicina Veterinária. Salvador (BA) – Brasil, 40pp. Teixeira, R.H.F., Amorim, M., Gazêta, G.S. & Serra-Freire, N.M. (2003) Ixodofauna de répteis cativos no Zoológico de Sorocaba, São Paulo, Brasil. Entomologia y Vectores, 10, (3), 319–329. Venzal, J.M., Autino, A.G., Nava, S. & Guglielmone, A.A. (2004) Ornithodoros mimon Kohls, Clifford and Jones, 1969 (Acari: Argasidae) on Argentinean bats, and new records from Uruguay. Systematic and Applied Acarology, 9, (1), 37–39. http://dx.doi.org/10.11158/saa.9.1.7 Venzal, J.M., Estrada-Peña, A., Mangold, A.J., González-Acuña, D. & Guglielmone, A.A. (2008) The Ornithodoros (Alectorobius) talaje species group (Acari: Ixodida: Argasidae): description of Ornithodoros (Alectorobius) rioplatensis n.sp. from southern South America. Journal of Medical Entomology, 45, 832–840. http://dx.doi.org/10.1603/0022-2585(2008)45[832:toatsg]2.0.co;2 Venzal, J.M., Nava, S., Mangold, A.J., Mastropaolo, M., Casás, G. & Guglielmone, A.A. (2012) Ornithodoros quilinensis sp. nov. (Acari, Argasidae), a new tick species from the Chacoan region in Argentina. Acta Parasitologica, 57, 329–336. http://dx.doi.org/10.2478/s11686-012-0034-5 Venzal, J.M., Nava, S., González-Acuña, D., Mangold, A.J., Muñoz Leal, S., Lado, P. & Guglielmone, A.A. (2013) A new species of Ornithodoros (Acari: Argasidae), parasite of Microlophus spp. (Reptilia: Tropiduridae) from northern Chile. Ticks and Tick-borne Diseases, 4, 128–132. http://dx.doi.org/10.1016/j.ttbdis.2012.10.038 Venzal, J.M., González-Acuña, D., Muñoz-Leal, S., Mangold, A. & Nava, S. (2015) Two new species of Ornithodoros (Ixodida: Argasidae) from the Southern Cone of South America. Experimental and Applied Acarology, 66, 127–139. https://doi.org/10.1007/s10493-015-9883-6 Viana, L.A., Winck, G.R., Almeida-Santos, M., Telles, F.B.S., Gazêta, G.S. & Rocha, C.F.D. (2012) New host records for Amblyomma rotundatum (Acari: Ixodidae) from Grussaí restinga, Rio de Janeiro, Brazil. Revista Brasileira de Parasitologia Veterinária, 21, (3), 319–322. https://doi.org/10.1590/S1984-29612012000300028 Witter, R., Martins, T.F., Campos, A.K., Melo, A.L.T., Corrêa, S.H.R., Morgado, T.O., Wolf, R.W., May-Júnior, J.A., Sinkoc, A.L., Strüssmann, C., Aguiar, D.M., Ross, R.V., Semedo, T.B.F., Campos, Z., Desbiez, A.L.J., Labruna, M.B. & Pacheco, R.C. (2016) Rickettsial infection in ticks (Acari: Ixodidae) of wild animals in midwestern Brazil. Ticks and Tick-borne Diseases, 7, (3), 415–423. https://doi.org/10.1016/j.ttbdis.2015.12.019 Woehl, G. (2002) Infestação de Amblyomma rotundatum (Koch) (Acari, Ixodidae) em sapos Bufo ictericus (Spix) (Amphibia, Bufonidae): novo registro de hospedeiro. Revista Brasileira de Zoologia, 19, 329–333. https://doi.org/10.1590/S0101-81752002000200003 Wolf, R.F., Aragona, M., Muñoz-Leal, S., Pinto, L.B., Melo, A.L.T. & Braga, I.A. (2016) Novel Babesia and Hepatozoon agents infecting non-volant small mammals in the Brazilian Pantanal, with the first record of the ticks Ornithodoros guaporensis in Brazil. Ticks and Tick-borne Diseases, 7, (3), 449–456. http://dx.doi.org/10.1016/j.ttbdis.2016.01.005 Submitted: 13 Jun. 2017; accepted by Trevor Petney: 24 Jun. 2018; published: 20 Jul. 2018

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