**Abstract**

Ticks are important ectoparasites and can transmit a wide variety of pathogens to animals and humans worldwide. These ectoparasites are the most important vectors of diseases causing pathogens in domestic and wild animals, and the second for man. In Brazil, Spotted Fever is the only disease transmitted to humans by ticks, while for animals we can highlight babesiosis, ehrlichiosis and anaplasmosis. Although ticks are found in all of Brazil's biomes, little is known about its diversity and host-relations in the Amazon biome. The existing gaps and the lack of research indicate that the diversity of ticks and their possible pathogens are underestimated in the Brazilian Amazon. Therefore, in order to guide the next studies in the Amazon biome, we present in this chapter a compilation of the records of hard ticks and soft ticks parasitizing wild and domestic animals, and humans. We present the general list of ticks for this biome, their distribution, hosts and importance for public health and veterinary. Finally, the ixodofauna found in the Brazilian Amazon and presented here does not match the vertebrate diversity of this biome, which is one of the largest on the planet. Therefore, more acarologists and epidemiologists are needed in this region.

**Keywords:** Ixodida, domestic animals, wild animals, humans, Amazon, Brazil

#### **1. Introduction**

The Amazon or Amazon Rainforest is the largest remnant of tropical forest in the world, occupying a region of approximately 6.7 million/km<sup>2</sup> , covering nine countries in South America: Ecuador (ffi 2%), Suriname (ffi 2%), Bolivia (ffi 4%), Venezuela (ffi 4%), Guyana (ffi 3%), French Guiana (ffi 2%), Colombia (ffi 10%), Peru (ffi 13%), and Brazil (ffi 60%) (**Figure 1**) [1, 2]. In Brazil, the Amazon biome, also known as "Legal Amazon" occupies approximately 49% of its territory, covering the states of Acre, Amapá, Amazonas, Mato Grosso, Pará, Rondônia, Roraima, Tocantins and Maranhão (**Figure 1**). The Brazilian Amazon is known for its high richness of landscapes composed of 23 ecoregions, whose main domain is the humid rainforest (ffi 78%). Due to this heterogeneity, the Amazon has an immeasurable amount of essential habitats for the maintenance of flora and fauna, represented

worldwide [8], therefore a concern for public health professional. In Brazil, ticks are vectors of diseases such as babesiosis, ehrlichiosis, anaplasmosis and rickettsioses,

Ticks belong to the Sub-Class Acari, Super-Order Parasitiformes, Order Ixodida, and four families: Ixodidae, Argasidae, Nuttallielidae and Deinocrotonidae [11, 12], the latter extinct. Of these, only the Ixodidae and Argasidae families occur in Brazil, with nine genera and 75 species [11, 13–16]. Ixodidae family is the most diverse with

51 species and five genera: *Amblyomma* (33 species), *Ixodes* (12 species),

*Ticks from the Brazilian Amazon: Species, Distribution and Host-Relations*

species). Argasidae family is represented by 24 species and four genera: *Ornithodoros* (18 species), *Antricola* (three species), *Argas* (one species) and *Nothoaspis* (two species). In general, the immature stages (larvae and nymphs) of two or three host ticks feed on small mammals and birds, while adults exploit medium to large-sized hosts [17]. One-host species *Dermacentor nitens* and *Rhipicephalus microplus*, complete the life cycle mainly on large animals such as horses and cattle, respectively. Second Esser et al. [18] the diversity of ticks increases with the increase of diversity of hosts, inasmuch as these hematophagous arthropods rely on the hosts to complete their life cycles. Therefore, if we take into

account the high diversity of hosts living in this biome and the amount of unexplored ecoregions, it is reasonable to infer that the diversity of ticks in the Amazon biome is insufficiently addressed. Thus, a review on the subject is necessary to point out the existing gaps and encourage new studies on ticks in the

The class Amphibia includes the orders Anura, Caudata, Gymnophiona, whereas the class Reptilia includes the orders Squamata, Testudines and Crocodylia. There are 331 amphibian and 550 reptile species in the Brazilian Amazon (**Figure 1**), although that faunal records are far from complete [2, 19]. Brazil has witnessed an increasing number of reports on tick parasitism of amphibians and reptiles over the past few years [20–24]. However, knowledge of this tick fauna as their hosts remains incomplete. To date, major tick-amphibian associations reported in Brazil are the ixodids *Amblyomma dissimile*, *Amblyomma rotundatum*, *Amblyomma humerale*, *Amblyomma fuscum* and *Amblyomma goeldii* [20–24], and the argasids *Ornithodoros saraivai* and *Ornithodoros faccinii* associated with species *Cycloramphus boraceiensis* and *Thoropa miliaris*, respectively [25, 26]. Occasional records on reptiles have also been reported for *Ornithodoros mimon* and *Ornithodoros rietcorreai* [22]*.* However, none of these argasids have been reported in the Amazon biome. Although the Brazilian Amazon has a high diversity of amphibians and reptiles, studies on ticks in association with these hosts are still rare, with a dominance of

The tick *A. dissimile* is constantly misidentified with the morphologically similar

reorganized, showing that this tick is restricted to the Pantanal and Amazon biomes [28]. Therefore, reports outside these biomes are considered misidentifications and must be viewed with caution. Most publications on this species are simple records of occurrence with some authors including data on prevalence and intensity of infestation both in the Amazon biome and elsewhere. According to Luz et al. [29] 12 (54.5%) out of 22 *Rhinella marina* toads captured in Amapá state, were parasitized by a total of 97 ticks (6 males, 39 females, 31 nymphs, 21 larvae) and mean intensity of 8.1 ticks per infested toad. In the Amazonian biome, *A. dissimile* is common on *R. marina* (Anura) and Boidae (Squamata) (**Table 1**). The experimental life cycle

**2. Hard ticks associated with amphibians and reptiles**

species *A. dissimile*, *A. rotundatum* and *A. humerale* [21–23, 27, 28].

*A. rotundatum*. Recently, the distribution of *A. dissimile* in the country was

*Rhipicephalus* (two species), *Haemaphysalis* (three species) and *Dermacentor* (one

including Brazilian Spotted Fever (BSF) [9, 10].

*DOI: http://dx.doi.org/10.5772/intechopen.94862*

Brazilian Amazon.

**35**

**Figure 1.**

*Diversity of reptiles, amphibians, birds and mammals in Brazil, including the Brazilian Amazon.*

with high biological diversity, which is regarded as the largest in the world [3, 4] The Amazon has approximately 45,000 species of flora (39,474 species) and fauna (5,526 species) [2, 5]. However, even with this expressive diversity and the numerous faunal studies carried out in the region, there are still many gaps in the biological diversity of the Amazon. Due to its enormous extension and high degree of preservation (e.g., unexplored areas), new bioecological associations and new species are discovered every year. In the past 20 years, more than 1,200 new species have been described in the Amazon region from which we can highlight 16 birds, 39 mammals, 55 reptiles and ffi 100 amphibians [2]. However, anthropic action has negatively impacted the animal-forest relationships for decades, resulting in the extinction of ecologically demanding species and, at the same time, the appearance of opportunistic and/or generalist species [6].

Many domestic and wild animals are responsible for the maintenance and dispersion of ticks in nature. In addition, vertebrates act as amplifiers and/or reservoirs for viruses, protozoa and bacteria transmitted by these ectoparasites. The antropic action affects the population dynamics of both ticks and their wild hosts directly or indirectly, and consequently, the epidemiology of tick-borne diseases once restricted to wild fauna can reach domestic animal and humans interfaces [7]. Ticks are responsible for more than 100,000 cases of diseases in humans and animals

*Ticks from the Brazilian Amazon: Species, Distribution and Host-Relations DOI: http://dx.doi.org/10.5772/intechopen.94862*

worldwide [8], therefore a concern for public health professional. In Brazil, ticks are vectors of diseases such as babesiosis, ehrlichiosis, anaplasmosis and rickettsioses, including Brazilian Spotted Fever (BSF) [9, 10].

Ticks belong to the Sub-Class Acari, Super-Order Parasitiformes, Order Ixodida, and four families: Ixodidae, Argasidae, Nuttallielidae and Deinocrotonidae [11, 12], the latter extinct. Of these, only the Ixodidae and Argasidae families occur in Brazil, with nine genera and 75 species [11, 13–16]. Ixodidae family is the most diverse with 51 species and five genera: *Amblyomma* (33 species), *Ixodes* (12 species), *Rhipicephalus* (two species), *Haemaphysalis* (three species) and *Dermacentor* (one species). Argasidae family is represented by 24 species and four genera: *Ornithodoros* (18 species), *Antricola* (three species), *Argas* (one species) and *Nothoaspis* (two species). In general, the immature stages (larvae and nymphs) of two or three host ticks feed on small mammals and birds, while adults exploit medium to large-sized hosts [17]. One-host species *Dermacentor nitens* and *Rhipicephalus microplus*, complete the life cycle mainly on large animals such as horses and cattle, respectively. Second Esser et al. [18] the diversity of ticks increases with the increase of diversity of hosts, inasmuch as these hematophagous arthropods rely on the hosts to complete their life cycles. Therefore, if we take into account the high diversity of hosts living in this biome and the amount of unexplored ecoregions, it is reasonable to infer that the diversity of ticks in the Amazon biome is insufficiently addressed. Thus, a review on the subject is necessary to point out the existing gaps and encourage new studies on ticks in the Brazilian Amazon.

### **2. Hard ticks associated with amphibians and reptiles**

The class Amphibia includes the orders Anura, Caudata, Gymnophiona, whereas the class Reptilia includes the orders Squamata, Testudines and Crocodylia. There are 331 amphibian and 550 reptile species in the Brazilian Amazon (**Figure 1**), although that faunal records are far from complete [2, 19]. Brazil has witnessed an increasing number of reports on tick parasitism of amphibians and reptiles over the past few years [20–24]. However, knowledge of this tick fauna as their hosts remains incomplete. To date, major tick-amphibian associations reported in Brazil are the ixodids *Amblyomma dissimile*, *Amblyomma rotundatum*, *Amblyomma humerale*, *Amblyomma fuscum* and *Amblyomma goeldii* [20–24], and the argasids *Ornithodoros saraivai* and *Ornithodoros faccinii* associated with species *Cycloramphus boraceiensis* and *Thoropa miliaris*, respectively [25, 26]. Occasional records on reptiles have also been reported for *Ornithodoros mimon* and *Ornithodoros rietcorreai* [22]*.* However, none of these argasids have been reported in the Amazon biome. Although the Brazilian Amazon has a high diversity of amphibians and reptiles, studies on ticks in association with these hosts are still rare, with a dominance of species *A. dissimile*, *A. rotundatum* and *A. humerale* [21–23, 27, 28].

The tick *A. dissimile* is constantly misidentified with the morphologically similar *A. rotundatum*. Recently, the distribution of *A. dissimile* in the country was reorganized, showing that this tick is restricted to the Pantanal and Amazon biomes [28]. Therefore, reports outside these biomes are considered misidentifications and must be viewed with caution. Most publications on this species are simple records of occurrence with some authors including data on prevalence and intensity of infestation both in the Amazon biome and elsewhere. According to Luz et al. [29] 12 (54.5%) out of 22 *Rhinella marina* toads captured in Amapá state, were parasitized by a total of 97 ticks (6 males, 39 females, 31 nymphs, 21 larvae) and mean intensity of 8.1 ticks per infested toad. In the Amazonian biome, *A. dissimile* is common on *R. marina* (Anura) and Boidae (Squamata) (**Table 1**). The experimental life cycle

with high biological diversity, which is regarded as the largest in the world [3, 4] The Amazon has approximately 45,000 species of flora (39,474 species) and fauna (5,526 species) [2, 5]. However, even with this expressive diversity and the numerous faunal studies carried out in the region, there are still many gaps in the biological diversity of the Amazon. Due to its enormous extension and high degree of preservation (e.g., unexplored areas), new bioecological associations and new species are discovered every year. In the past 20 years, more than 1,200 new species have been described in the Amazon region from which we can highlight 16 birds, 39 mammals, 55 reptiles and ffi 100 amphibians [2]. However, anthropic action has negatively impacted the animal-forest relationships for decades, resulting in the extinction of ecologically demanding species and, at the same time, the appearance

*Diversity of reptiles, amphibians, birds and mammals in Brazil, including the Brazilian Amazon.*

Many domestic and wild animals are responsible for the maintenance and dispersion of ticks in nature. In addition, vertebrates act as amplifiers and/or reservoirs for viruses, protozoa and bacteria transmitted by these ectoparasites. The antropic action affects the population dynamics of both ticks and their wild hosts directly or

restricted to wild fauna can reach domestic animal and humans interfaces [7]. Ticks are responsible for more than 100,000 cases of diseases in humans and animals

indirectly, and consequently, the epidemiology of tick-borne diseases once

of opportunistic and/or generalist species [6].

**Figure 1.**

*Ecosystem and Biodiversity of Amazonia*

**34**


including pre-attachment periods for each parasitic stage lasts approximately 350 days, as reported by Schumaker et al. [43] who started a colony from one engorged female collected from the Amazonian biome. Ogrzewalska et al. [37] reported *Rickettsia bellii* and '*Candidatus* Rickettsia colombianensi' in ticks collected from *Bothrops atrox* from Pará state. Luz et al. [29] reported '*Ca.* R. colombianensi'

*Hosts of* Amblyomma dissimile *and* Amblyomma rotundatum *in the Brazilian Amazon.*

**Host Specie** *A. dissimile A. rotundatum*

*Ticks from the Brazilian Amazon: Species, Distribution and Host-Relations*

*DOI: http://dx.doi.org/10.5772/intechopen.94862*

Iguanidae *Iguana iguana* 31 118M;

Total 4 2189 983M;

**Table 1.**

**37**

Chelidae *Platemys platycephala* 5F [32] Tropiduridade *Uranoscodon superciliosus* 7 6F [33]

Teiidae *Ameiva ameiva* 4 1F [31, 34]

Dactyoloidae *Norops auratus* 1 [34] Gekkonidae *Hemidactylus mabouia* 1 [34] Alligatoridae *Paleosuchus trigonatus* 9M; 15F 4 12F [31, 32]

**L N A LN A**

*Tropidurus* sp. 1M [38]

69F

*Caiman crocodilus* 3 [31, 39]

1496F

221 307F

2F [31, 36, 39, 40]

The tick *A. rotundatum* is an obligate parthenogenetic species, although there are six reports of adult males, two males in the laboratory [44] and six in the Amazon region parasitizing *Tropidurus* sp., *Boa constrictor*, *Iguana iguana*, *Chelonoidis denticulatus* and *B. atrox* [30, 36, 44–47]. Most publications on *A. rotundatum* are simple records of its occurrence with some additional data on prevalence and intensity of infestation in the Cerrado, Atlantic forest and Amazon biomes [22, 23, 31, 34, 48]. In the Brazilian Amazon, the families Bufonidae (Amphibian) and Boidae (Reptilia) are the most frequently parasitized by *A. rotundatum*, but there are reports on other species of Amphibian and Reptiles (**Table 1**). Recently, Gianizella et al. [31] reported *A. rotundatum* in different municipalities of the Amazonas state, on *B. constrictor* and several unknown hosts. The bioecology of this species under quasi-natural environment demonstrated a peak of larvae and nymphs in the dry season and females in the rainy season; however, there is a hypothesis of the absence of seasonality in this tick [48, 49]. Although *A. rotundatum* is a three-host tick on amphibians, it can behave as a two-host tick when feeding on snakes [50]. The complete experimental life cycle, including preattachment periods for each parasitic stage, ranged from 126 to 228 days on toads [51] or 56 to 163 days on snakes [50] and ulcerative lesions and hemorrhages after *A. rotundatum* feeding [48] were reported for ticks collected on toads from the Cerrado biome. Transmission of the hemogregarine *Hemolivia stellata* by *R. marina* collected in Belém, state of Pará was reported by Petit et al. [52]. This tick has also been found infected with *R. bellii* in the state of Amazon in the municipalities of Cacaulândia and Monte Negro, in the state of Rondônia; in the municipalities of Amapá (Ilha de Maracá) and Santana, in the state of Amapá; and in the municipality

in ticks collected on *R. marina* from Amapá state.

of Rio Branco, in the state of Acre [29, 53, 54].


*Ticks from the Brazilian Amazon: Species, Distribution and Host-Relations DOI: http://dx.doi.org/10.5772/intechopen.94862*

#### **Table 1.**

**Host Specie** *A. dissimile A. rotundatum*

Bufonidae *Rhinella marina* 37 12M; 16F 164 133F [23, 29–33]

Amphibian (Anura)

Reptile

**36**

Leptodactylidae *Leptodactylus*

*Ecosystem and Biodiversity of Amazonia*

*pentadactylus*

Boidae *Boa constrictor* 1703 723M;

*Erythrolamprus reginae semilineatus*

Viperidae *Bothrops atrox* 4 11 24M; 14F 1M;

Elapidae *Micrurus averyi* 3F [31]

Colubridade *Leptophis ahaetulla* 3 [33]

Dipsadidae *Xenodon severus* 8 [40]

Podocnemididae *Podocnemis expansa* 1F [39]

Kinosternidae *Kinosternon scorpioides* 2F [42]

*Chelonoidis carbonaria* 1 1M; 1F 11 [32, 39]

*Podocnemis unifilis* 1 [39] *Trachemys dorbigni* 2 12F [39]

Testudinidae *Chelonoidis denticulatus* 3M 3 2M;

**L N A LN A**

*Rhinella major* 2F 15 2F [33–35] *Rhaebo guttatus* 1 1F [23] *Rhinella margaritifera* 3 1F [23] *Rhinella gildae* 1 [36]

1346F

*Corallus hortulanus* 3 1M [31, 34] *Eunectes murinus* 305 21M; 3F [31, 40]

*Crotalus durissus* 1M; 1F [34] *Lachesis muta* 1 [34]

*Micrurus lemniscatus* 12M; 9F [31]

*Chironius multiventris* 3 [33] *Chironius scurrulus* 1M; 1F [34] *Chironius laevicollis* 12 15M; 6F [31] *Mastigodryas boddaerti* 1 [33] *Hydrodynastes gigas* 1 7F [30] *Helicops polylepis* 1 [34] *Leptodeira annulata* 1 [34] *Phimophis guerini* 1F [34] *Erythrolamprus reginae* 1 [34] *Spilotes pullatus* 72 42M; 9F [31]

1F [23]

[23, 30, 32– 34, 36–40]

[32, 33, 36, 37]

12 1M; 77F

2F

1 [34]

42F

[31, 41]

*Hosts of* Amblyomma dissimile *and* Amblyomma rotundatum *in the Brazilian Amazon.*

including pre-attachment periods for each parasitic stage lasts approximately 350 days, as reported by Schumaker et al. [43] who started a colony from one engorged female collected from the Amazonian biome. Ogrzewalska et al. [37] reported *Rickettsia bellii* and '*Candidatus* Rickettsia colombianensi' in ticks collected from *Bothrops atrox* from Pará state. Luz et al. [29] reported '*Ca.* R. colombianensi' in ticks collected on *R. marina* from Amapá state.

The tick *A. rotundatum* is an obligate parthenogenetic species, although there are six reports of adult males, two males in the laboratory [44] and six in the Amazon region parasitizing *Tropidurus* sp., *Boa constrictor*, *Iguana iguana*, *Chelonoidis denticulatus* and *B. atrox* [30, 36, 44–47]. Most publications on *A. rotundatum* are simple records of its occurrence with some additional data on prevalence and intensity of infestation in the Cerrado, Atlantic forest and Amazon biomes [22, 23, 31, 34, 48]. In the Brazilian Amazon, the families Bufonidae (Amphibian) and Boidae (Reptilia) are the most frequently parasitized by *A. rotundatum*, but there are reports on other species of Amphibian and Reptiles (**Table 1**). Recently, Gianizella et al. [31] reported *A. rotundatum* in different municipalities of the Amazonas state, on *B. constrictor* and several unknown hosts. The bioecology of this species under quasi-natural environment demonstrated a peak of larvae and nymphs in the dry season and females in the rainy season; however, there is a hypothesis of the absence of seasonality in this tick [48, 49]. Although *A. rotundatum* is a three-host tick on amphibians, it can behave as a two-host tick when feeding on snakes [50]. The complete experimental life cycle, including preattachment periods for each parasitic stage, ranged from 126 to 228 days on toads [51] or 56 to 163 days on snakes [50] and ulcerative lesions and hemorrhages after *A. rotundatum* feeding [48] were reported for ticks collected on toads from the Cerrado biome. Transmission of the hemogregarine *Hemolivia stellata* by *R. marina* collected in Belém, state of Pará was reported by Petit et al. [52]. This tick has also been found infected with *R. bellii* in the state of Amazon in the municipalities of Cacaulândia and Monte Negro, in the state of Rondônia; in the municipalities of Amapá (Ilha de Maracá) and Santana, in the state of Amapá; and in the municipality of Rio Branco, in the state of Acre [29, 53, 54].

Most reports of *A. humerale* mention the adult stage parasitizing species of tortoises, namely the "yellow-footed tortoise" *C. denticulatus* and the "red-footed tortoise" *Chelonoidis carbonarius* [27, 55]. There is also a record of *A. humerale* on *Rhinoclemmys punctularia* in central Amazon [56]. Labruna et al. [27] collected 215 adult ticks from six *C. denticulatus* and nine *Chelonoidis* sp. from state of Rondônia, with mean infestation of 14.3 12.0 ticks per tortoise. In addition, seven engorged nymphs were collected on lizards. Morais et al. [57] collected 120 adult ticks from 18 (75%) out of 24 *C. denticulatus* captured in a transitional area between the biomes Cerrado and southwestern Amazon rainforest, in the state of Mato Grosso. The mean intensity of infestation was 6.7 ticks/tortoise. In both surveys, male ticks were mostly attached in clusters on the carapace whereas females were found attached to the tortoise skin. The sex ratio (males:females) for *A. humerale* were different in both surveys, 10.3:1 [27] and 1.1:1 [57]. There is one additional record of two nymphs on *Paleosuchus trigonatus* (Crocodylia) [32]. The life-cycle in the laboratory, including pre-feeding periods for each of the parasitic stages, could be completed in an average period of ca. 200 days [58]. *Rickettsia bellii* and *Rickettsia amblyommatis* have been detected in *A. humerale* [42, 53]. The records of *A. fuscum* in association with amphibians and reptiles are rare in Brazil. Dantas-Torres et al. [59] collected one male tick in one out of 490 caimans (*Caiman latirostris* and *Paleosuchus palpebrosus*) trapped in the Atlantic rainforest biome in Pernambuco state, northeastern Brazil. *Amblyomma goeldii* has been recorded only in the Amazonas state. Martins et al. [60] reported two males collected on *B. constrictor* and recovery of ≈100 (20%) engorged larvae out of ≈500 unfed larvae experimentally infested on a *B. constrictor.*

of Brazilian Amazonian have been recorded in association with at least one tick species (**Table 2**). This is equivalent to approximately only 7% of bird species found in this biome and 5% of the total birds recorded in Brazil (**Table 2**). Similar to other studies regarding tick-bird associations in Brazil [73–75], Passeriformes birds were the most parasitized in the Amazon biome, including 14 families and 72 species (**Figure 2** and **Table 2**). In this group, the greatest diversity of parasitized birds was Thamnophilidae (20 species) followed by Dendrocolaptidae (16 species) and Tyrannidade (10 species) (**Figure 2**). The least parasitized families were

*Ticks from the Brazilian Amazon: Species, Distribution and Host-Relations*

*DOI: http://dx.doi.org/10.5772/intechopen.94862*

Conopophagidae, Furnariidae, Xenopidae, Tityridae, Cardinalidae, Columbidae, Cuculidae, Momotidae, Capitonidae, Ramphastidae, Psittacidae, Accipitridae and Falconidae with only one species of parasitized bird each (**Figure 2** and **Table 2**). Non-Passerines were represented by 10 different orders and 11 families, with

**Hosts Ticks References**

*schistogynus*

*Thamnophilus schistaceus*

*Myrmelastes hyperythrus*

*Sciaphylax hemimelaena*

*Schistocichla leucostigma*

*Phlegopsis nigromaculata*

*Hylophylax punctulatus*

*Hylophylax poecilinotus*

**39**

*Thamnophilus aethiops A. calcaratum*

*Hylophylax naevius H. juxtakochi*

*Thamnomanes caesius A. geayi*

Passeriformes Thamnophilidae *Thamnomanes*

**Species Stages**

*A. humerale A. nodosum*

*Thamnophilus doliatus A. nodosum* 2 [68]

*A. nodosum Amblyomma* sp.

*A. longirostre*

*A. humerale Amblyomma* sp.

*A.coelebs A. longirostre A. nodosum Amblyomma* sp.

*Hypocnemis subflava Amblyomma* sp. 3 [70]

*A. humerale A. longirostre A. calcaratum Amblyomma* sp.

*A. longirostre A. humerale Amblyomma* sp

*A. geayi A. humerale A. longirostre Amblyomma* sp. *H. juxtakochi*

*Myrmornis torquata Amblyomma* sp. 3 [69]

LL NN

1 1

5 1

*A. nodosum* 9 [68]

*A. nodosum* 1 [68]

4 5

1 1 24

*Amblyomma* sp. 1 [69]

1 3 4

1 2 1

2 1

2 2 [68]

[68]

[68]

[69]

[69]

[69, 71]

[69]

5 [68, 69]
