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

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 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


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

The importance of birds to maintain biodiversity and ecological balance of nature is notorious [61]. Due to migration, wild birds are of concern to human and animal health worldwide [62] because they can carry infected ticks over long distances, directly influencing the epidemiology of tick-borne diseases in animals and humans. In addition, wild birds themselves can be reservoirs of *Borrelia burgdorferi* sensu lato, and potentially to *Anaplasma phagocytophilum* and *Rickettsia* spp. [62, 63]. Wild birds play an important role in maintaining and dispersing immatures (larvae and nymphs) of several tick species into new locations [61, 64]. Of the total genera of ticks described in Brazil, five have at least one species recorded in association with wild birds. The most common are the hard ticks of the genera *Amblyomma*, *Haemaphysalis* and *Ixodes* [61, 64]. There are also occasional

reports of the genera *Rhipicephalus* and *Ornithodoros*. Ticks of the genus

BSF by dispersing and maintaining their vectors in nature.

*Amblyomma* are the most common on wild birds in the Brazil including the Amazon biome, exclusively for the larvae and nymphs [61, 64]. Adult ticks are only occasionally found on wild birds, with the exception of *Ixodes paranaensis* and *Ixodes auritulus*, which have the entire cycle synchronized with birds [65]. In Brazil, there are no reports of wild birds as a source of pathogens transmitted by ticks to humans, but they can serve as disperser hosts for vectors of Brazilian Spotted Fever (BSF) as *Amblyomma sculptum*, *Amblyomma aureolatum* and *Amblyomma ovale*, in the larvae and nymph stages [64]. Therefore, wild birds act indirectly in the epidemiology of

Over more than 1,900 birds recorded in Brazil, approximately 1,300 reside in the Brazilian Amazon, with a 20% of endemism [66] (**Figure 1**). Of these, approximately 7% are migrants from the northern hemisphere and southern South America, including migrations from other Brazilian biomes [67]. To date, 86 bird species

*B. constrictor.*

**38**

**3. Hard ticks associated with wild birds**

*Ecosystem and Biodiversity of Amazonia*


**Hosts Ticks References**

*Xiphorhynchus ocellatus*

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

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

*Xiphorhynchus pardalotus*

Pipridae *Pipra filicauda A. geayi*

*Machaeropterus pyrocephalus*

*amaurocephalus*

*Todirostrum maculatum*

*Hemitriccus flammulatus*

*Onychorhynchus coronatus*

*Ramphotrigon megacephalum*

*Knipolegus poecilocercus*

**41**

Rhynchocyclidae *Leptopogon*

*Dendroplex picus A. longirostre*

*Pipra fasciicauda A. longirostre*

*Dixiphia pipra A. geayi*

*Rhynchocyclus olivaceus A. geayi*

*Poecilotriccus latirostris A. nodosum*

Tyrannidae *Myiozetetes similis A. nodosum* 1 [68] *Empidonax alnorum A. nodosum*

*Mionectes macconnelli A. longirostre*

*Attila spadiceus A. geayi*

Xenopidae *Xenops minutus A. nodosum* 2 [69]

**Species Stages**

*A. nodosum*

*Amblyomma* sp.

*A. nodosum Amblyomma* sp.

*A. longirostre*

*A. humerale A. longirostre A. nodosum Amblyomma* sp.

*Amblyomma* sp.

*A. longirostre*

*Amblyomma* sp.

*Amblyomma* sp.

*Myiobius barbatus A. longirostre* 3 [69]

*A. longirostre Amblyomma* sp.

*A. nodosum Amblyomma* sp.

*Platyrinchus saturatus Amblyomma* sp. 1 [69] *Hemitriccus minor A. longirostre* 1 [69]

*Lepidothrix nattereri Amblyomma* sp. 1 [71]

*A. calcaratum* 1 [71]

*Amblyomma* sp. 3 [31]

1 8

*A. longirostre* 1 [68]

1 1

*A. geayi* 1 [68]

1 1 4

*A. nodosum* 1 [68]

*Amblyomma* sp. 1 [70]

4 5

1 3

3 6

*Amblyomma* sp. 1 [71]

1 1

2 1 17

6 3 [70]

4 1 [68]

[69]

[69]

[69]

1 3 [70]

3 3 [68, 70]

1 [68, 70]

7 [68]

[69]

[68]


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

**Hosts Ticks References**

*Myrmotherula hauxwelli*

*Ecosystem and Biodiversity of Amazonia*

*Myrmotherula longipennis*

*Myrmoborus myotherinus*

*Epinecrophylla leucophthalma*

Dendrocolaptidae *Dendrocincla merula A. longirostre*

*Deconychura longicauda*

*Deconychura stictolaema*

*Dendrocolaptes hoffmannsi*

*Sittasomus griseicapillus*

*Xiphorhynchus guttatoides*

**40**

*Glyphorynchus spirurus A. geayi*

*Hylexetastes brigidai A. geayi*

*Xiphorhynchus guttatus A. longirostre*

*Xiphorhynchus elegans A. longirostre*

*Pyriglena leuconota A. coelebs*

Conopophagidae *Conopophaga aurita Amblyomma* sp. 1 [69] Scleruridae *Sclerurus cf. rufigularis A. humerale* 1 [68]

Furnariidae *Automolus paraensis Amblyomma* sp. 1 [69]

**Species Stages**

*A. longirostre Amblyomma* sp.

*A. humerale A. longirostre Amblyomma* sp.

*Willisornis poecilinotus A. calcaratum* 1 [71]

*Sclerurus caudacutus A. humerale* 1 [69]

*Amblyomma* sp.

*Dendrocincla fuliginosa Amblyomma* sp. 10 [31, 70]

*A. longirostre Amblyomma* sp.

*A. longirostre Amblyomma* sp.

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

*Dendrocolaptes certhia A. longirostre* 1 [71] *Glyphorynchus spirurus A. longirostre* 1 [71]

> *A. longirostre Amblyomma* sp.

> *A. longirostre Amblyomma* sp.

> *A. humerale Amblyomma* sp.

> > *A. nodosum*

*A. humerale Amblyomma* sp.

*Amblyomma* sp.

*Amblyomma* sp. 1 [69]

3 3

*A. longirostre* 1 [69]

*A. longirostre* 1 [71]

7 34

4 39

> 1 1

1 6 16

1 6 21

> 3 2

2 1 3

1 2 [69]

1 [69]

1 [68, 69, 71]

1 [69, 71]

1 [69]

[69]

[69]

[68]

2 [69]

2 1 [70]

1 2 [70]

2 [69]

2 1


emphasis on Bucconidae with three species (**Figure 2** and **Table 2**). To date, approximately 1,068 specimens of ticks have been collected from birds in the Brazilian Amazon, in the stages of larvae (884/83%), nymphs (184/17%) and no adults (**Figure 3**). These are included in the genera *Amblyomma* and *Haemaphysalis*. The genus *Amblyomma* was the dominant with eight species (**Table 2**). The greatest diversity of ticks was reported for the Thamnophilidae family with seven species: *A. longirostre*, *A. nodosum*, *A. humerale*, *A. calcaratum*, *A. geayi*, *A. coelebs* and *Haemaphysalis juxtakochi*. The Dendrocolaptidae family was the second with five

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

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

**Figure 2.**

**Figure 3.**

**43**

*Diversity of hard ticks parasitizing wild birds from the Brazilian Amazon.*

*Distribution of studies reporting the association of hard ticks on wild birds from the Brazilian Amazon.*

#### **Table 2.**

*Ticks identified on wild birds in the Brazilian Amazon biome.*

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

emphasis on Bucconidae with three species (**Figure 2** and **Table 2**). To date, approximately 1,068 specimens of ticks have been collected from birds in the Brazilian Amazon, in the stages of larvae (884/83%), nymphs (184/17%) and no adults (**Figure 3**). These are included in the genera *Amblyomma* and *Haemaphysalis*. The genus *Amblyomma* was the dominant with eight species (**Table 2**). The greatest diversity of ticks was reported for the Thamnophilidae family with seven species: *A. longirostre*, *A. nodosum*, *A. humerale*, *A. calcaratum*, *A. geayi*, *A. coelebs* and *Haemaphysalis juxtakochi*. The Dendrocolaptidae family was the second with five

**Figure 2.**

**Hosts Ticks References**

*genibarbis*

*Microcerculus marginatus*

Tityridae *Schiffornis turdina A. longirostre*

*Turdus amaurochalinus*

Thraupidae *Ramphocelus carbo A. nodosum*

*Tachyphonus surinamus*

Columbiformes Columbidae *Columbina talpacoti A. nodosum* 1 [68] Cuculiformes Cuculidae *Crotophaga major A. nodosum* 1 [68]

*insperatus*

Galbuliformes Bucconidae *Monasa nigrifrons Amblyomma* sp. 1 [70]

Psittaciformes Psittacidae *Primolius maracana A. dissimile* 1 [72] Accipitriformes Accipitridae *Harpia harpyja A. cajennense* 16 [41]

Cariamiformes Cariamidae *Cariama cristata A. cajennense* 5 [39] Galliformes Cracidae *Penelope superciliaris A. cajennense* 1 1 [42] Falconiformes Falconidae *Micrastur ruficollis A. longirostre* 1 [71]

**Total 884 184**

Coraciiformes Momotidae *Momotus momota A. longirostre*

Piciformes Capitonidae *Capito auratus*

*Ticks identified on wild birds in the Brazilian Amazon biome.*

**Table 2.**

**42**

Troglodytidae *Pheugopedius*

*Ecosystem and Biodiversity of Amazonia*

**Species Stages**

*Amblyomma* sp. 3 [69]

3 14

*Amblyomma* sp. 1 [70]

2 3

1 1 5

44 15 4

> 1 1

2 1

*H. juxtakochi* 1 [41]

363 1 [68]

[69]

[68]

3 [31, 69]

[30, 68, 70, 71]

[68, 70]

[68]

*A. nodosum Amblyomma* sp.

*Amblyomma* sp.

*Turdus sanchezorum A. longirostre* 1 [68]

*A. nodosum A. longirostre Amblyomma* sp.

*A. coelebs Amblyomma* sp.

*A. longirostre Amblyomma* sp.

*A. humerale A. longirostre Amblyomma* sp.

*A. humerale*

*A. nodosum A. longirostre*

*Pteroglossus bitorquatus A. longirostre* 1 [30]

*Malacoptila rufa Amblyomma* sp. 1 [69] *Galbula cyanicollis Amblyomma* sp. 2 [69]

Turdidae *Turdus hauxwelli A. longirostre* 2 [68]

*Turdus ignobilis A. geayi*

*Turdus albicollis A. longirostre*

Cardinalidae *Cyanoloxia cyanoides Amblyomma* sp. 1 [71]

Ramphastidae *Ramphastos vitellinus A. geayi* 3 [31]

**Figure 3.** *Distribution of studies reporting the association of hard ticks on wild birds from the Brazilian Amazon.*

important role in preserved or anthropized ecosystems. Therefore, the knowledge of the local diversity of wild mammals, and their relationship with ticks is considered an important tool for public conservation policies and consequently for public health. The alteration of wild habitats can determine changes in the patterns of parasitic specificity, inducing tick species to seek new groups of hosts, increasing the risk of disease transmission [79, 80]. Some wild mammals (small, medium and large) are directly or indirectly involved in the transmission cycles of many tickborne pathogens worldwide, including *Anaplasma* spp., *Babesia* spp., *Borrelia* spp., and *Rickettsia* spp. [80, 81]. In Brazil, the main zoonosis transmitted by ticks is Brazilian Spotted Fever caused by the bacterium *Rickettsia rickettsii*, which has the rodent *Hydrochoerus hydrochaeris* as its main amplifier. In addition, there is evidence that marsupials and small rodents can serve as amplifier hosts for *R. rickettsii* in

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

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

In Brazil there are approximately 755 species of mammals distributed in all its six biomes, including the Amazon biome [78, 79]. Of the total mammal species, 41% (ffi 311 species) occur in the Amazon biome (**Figure 1**) [2, 76]. These vertebrates are distributed in 11 orders, 51 families and 249 genera [78, 79]. Among the families, Cricetidae is the most diverse, with 144 species [78, 79]. The vast majority of tick species, including all life stages, in Brazil have records on wild mammals of different sizes [17]. In general, small mammals of the orders Rodentia and Didelphimorphia are those that have a greater number of studies in association with ticks, especially the families Cricetidae and Didelphidae [17, 82]. In general, medium and large mammals are parasitized by ticks in all stages (larva, nymph and adult), while in small mammals the stages of larva and nymph are more common. In this last group of hosts we can highlight the cricetids *Akodon* spp., *Calomys* spp., *Oligoryzomys* spp. and *Nectomys* spp. as the most parasitized by immature ticks in

Nine orders and 24 families of wild mammals have representatives in association with ticks in the Brazilian Amazon (**Figure 5**). Rodentia was the most diverse with

Didelphimorphia (12 species) (**Figure 5**). However, it was the family Didelphidae that presented a greater number of parasitized species, followed by Dasyproctidae (seven species) and Mustelidae (four species). These records corroborate with numerous studies of tick parasitism on wild animals from South America, with

Hard ticks parasitizing wild mammals in the Brazilian Amazon are represented by five genera: *Amblyomma*, *Ixodes*, *Haemaphysalis*, *Dermacentor* and *Rhipicephalus*. Of these, the genus *Amblyomma* was more frequently recorded with 23 species (**Figure 5**). The orders Rodentia, Pilosa and Didelphimorphia are hosts for highest diversity to *Amblyomma* species 17, 15 and eight, respectively. All these species are

The second most common genus in the Amazon is *Ixodes*, with six reported species: *Ixodes amarali*, *Ixodes bocatorensis*, *Ixodes lasallei*, *Ixodes luciae*, *Ixodes schulzei* and *Ixodes spinosus* (**Table 3**)*.* Although there are two exclusive species on birds in Brazil [65], all species recorded in the Brazilian Amazon parasitize mainly wild mammals [13, 16, 31, 32, 88, 99, 102]. *Ixodes* spp. were found on families Didelphidae, Dasyproctidae, Cricetidae, Myrmecophagidae, Bradypodidae and Cyclopedidae (**Figure 5** and **Table 3**). In general *Ixodes* ticks mainly parasitize

16 species of mammals, followed by the orders Carnivora (13 species) and

emphasis on the orders Rodentia and Didelphimorphia [17, 82, 83].

also recorded on a variety of wild mammals in Brazil [83–87], except for *A. rotundatum* and *A. dissimile,* which are more specific ticks of cold-blooded animals (Amphibians and Reptiles), although there are occasional reports on mammals in South America [20, 83]. *Amblyomma humerale,* of which the adult stage is more specific to tortoises, immature stages have been found on a variety of

small mammals, reptiles and birds [17, 68, 71, 83].

nature [82].

nature.

**45**

**Figure 4.**

*Hard tick species parasitizing wild birds from the Brazilian Amazon.*

species: *A. longirostre*, *A. nodosum*, *A. humerale*, *A. calcaratum* and *A. geayi* (**Figure 2** and **Table 2**).

Overall, *A. longirostre* and *A. nodosum* are the two most common species on wild birds in the Brazilian Amazon [68–71] (**Figure 4** and **Table 2**). The tick *A. longirostre* was the most common, recorded in 12 families and 40 species of birds (36 Passerines and 4 non-Passerines) (**Figure 2**). *Amblyomma longirostre* has been treated as an arboreal tick, with immatures parasitizing birds and adults parasitizing rodents Erethizontidae (e.g., *Sphiggurus* spp.) [73, 76]. This tick was also the most abundant with 110 larvae and 39 nymphs. *Amblyomma longirostre* was found in co-infestation with the following species: *A. calcaratum*, *A. nodosum*, *A. coelebs*, *A. humerale*, *A. geayi* and *H. juxtakochi* (**Table 2**). In addition, as it is frequent on birds, *A. longirostre* is popularly known as "bird tick" or "bird earring" [77]. *Amblyomma nodosum* was the second most common species collected on 12 families and 22 species of birds. This tick was recorded in co-infestation with *A. longirostre*, *A. coelebs*, *A. humerale* and *A. geayi* (**Table 2** and **Figures 2** and **4**). The birds most infested by *A. nodosum* were *Rhynchocyclus olivaceus* and *Ramphocelus carbo* with 17 and 15 nymphs, respectively. These birds inhabit the forest understory and visit the soil occasionally [67]. As Xenartha mammals (*Myrmecophaga tridactyla*,*Tamandua tetradactyla*) are the primary hosts of *A. nodosum* [17], it is believed that the low areas of the understory are a major source of infestation. Interestingly, *T. tetradactyla* may have arboreal habits [78], which helps to explain the presence of *A. nodosum* also on birds of different forest strata.

To date, of the total of ticks collected, 736 (70%) were larvae identified as *Amblyomma* sp. due to lack of reliable tools for larval identification, thus, indicating that the diversity of ticks on birds of the Amazon may be underestimated. Luz et al. [75], using molecular biology, identified more than 90% of the larvae collected from birds in the Atlantic Forest biome, reporting the greatest diversity of ticks on birds in a single study in Brazil and description of the new tick *Amblyomma romarioi* [14]. Therefore, the identification of all larvae by molecular biology in addition to morphological identification is extremely important to ascertain the diversity of ticks in the Amazon biome.

#### **4. Hard ticks associated with wild and domestic mammals**

The fauna of wild mammals in Brazil is quite diverse and more than half lives in the Amazon biome [79]. Like birds, amphibians and reptiles, mammals play an

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

important role in preserved or anthropized ecosystems. Therefore, the knowledge of the local diversity of wild mammals, and their relationship with ticks is considered an important tool for public conservation policies and consequently for public health. The alteration of wild habitats can determine changes in the patterns of parasitic specificity, inducing tick species to seek new groups of hosts, increasing the risk of disease transmission [79, 80]. Some wild mammals (small, medium and large) are directly or indirectly involved in the transmission cycles of many tickborne pathogens worldwide, including *Anaplasma* spp., *Babesia* spp., *Borrelia* spp., and *Rickettsia* spp. [80, 81]. In Brazil, the main zoonosis transmitted by ticks is Brazilian Spotted Fever caused by the bacterium *Rickettsia rickettsii*, which has the rodent *Hydrochoerus hydrochaeris* as its main amplifier. In addition, there is evidence that marsupials and small rodents can serve as amplifier hosts for *R. rickettsii* in nature [82].

In Brazil there are approximately 755 species of mammals distributed in all its six biomes, including the Amazon biome [78, 79]. Of the total mammal species, 41% (ffi 311 species) occur in the Amazon biome (**Figure 1**) [2, 76]. These vertebrates are distributed in 11 orders, 51 families and 249 genera [78, 79]. Among the families, Cricetidae is the most diverse, with 144 species [78, 79]. The vast majority of tick species, including all life stages, in Brazil have records on wild mammals of different sizes [17]. In general, small mammals of the orders Rodentia and Didelphimorphia are those that have a greater number of studies in association with ticks, especially the families Cricetidae and Didelphidae [17, 82]. In general, medium and large mammals are parasitized by ticks in all stages (larva, nymph and adult), while in small mammals the stages of larva and nymph are more common. In this last group of hosts we can highlight the cricetids *Akodon* spp., *Calomys* spp., *Oligoryzomys* spp. and *Nectomys* spp. as the most parasitized by immature ticks in nature.

Nine orders and 24 families of wild mammals have representatives in association with ticks in the Brazilian Amazon (**Figure 5**). Rodentia was the most diverse with 16 species of mammals, followed by the orders Carnivora (13 species) and Didelphimorphia (12 species) (**Figure 5**). However, it was the family Didelphidae that presented a greater number of parasitized species, followed by Dasyproctidae (seven species) and Mustelidae (four species). These records corroborate with numerous studies of tick parasitism on wild animals from South America, with emphasis on the orders Rodentia and Didelphimorphia [17, 82, 83].

Hard ticks parasitizing wild mammals in the Brazilian Amazon are represented by five genera: *Amblyomma*, *Ixodes*, *Haemaphysalis*, *Dermacentor* and *Rhipicephalus*. Of these, the genus *Amblyomma* was more frequently recorded with 23 species (**Figure 5**). The orders Rodentia, Pilosa and Didelphimorphia are hosts for highest diversity to *Amblyomma* species 17, 15 and eight, respectively. All these species are also recorded on a variety of wild mammals in Brazil [83–87], except for *A. rotundatum* and *A. dissimile,* which are more specific ticks of cold-blooded animals (Amphibians and Reptiles), although there are occasional reports on mammals in South America [20, 83]. *Amblyomma humerale,* of which the adult stage is more specific to tortoises, immature stages have been found on a variety of small mammals, reptiles and birds [17, 68, 71, 83].

The second most common genus in the Amazon is *Ixodes*, with six reported species: *Ixodes amarali*, *Ixodes bocatorensis*, *Ixodes lasallei*, *Ixodes luciae*, *Ixodes schulzei* and *Ixodes spinosus* (**Table 3**)*.* Although there are two exclusive species on birds in Brazil [65], all species recorded in the Brazilian Amazon parasitize mainly wild mammals [13, 16, 31, 32, 88, 99, 102]. *Ixodes* spp. were found on families Didelphidae, Dasyproctidae, Cricetidae, Myrmecophagidae, Bradypodidae and Cyclopedidae (**Figure 5** and **Table 3**). In general *Ixodes* ticks mainly parasitize

species: *A. longirostre*, *A. nodosum*, *A. humerale*, *A. calcaratum* and *A. geayi*

*Hard tick species parasitizing wild birds from the Brazilian Amazon.*

Overall, *A. longirostre* and *A. nodosum* are the two most common species on wild birds in the Brazilian Amazon [68–71] (**Figure 4** and **Table 2**). The tick *A. longirostre* was the most common, recorded in 12 families and 40 species of birds (36 Passerines and 4 non-Passerines) (**Figure 2**). *Amblyomma longirostre* has been treated as an arboreal tick, with immatures parasitizing birds and adults parasitizing rodents Erethizontidae (e.g., *Sphiggurus* spp.) [73, 76]. This tick was also the most abundant with 110 larvae and 39 nymphs. *Amblyomma longirostre* was found in co-infestation with the following species: *A. calcaratum*, *A. nodosum*, *A. coelebs*, *A. humerale*, *A. geayi* and *H. juxtakochi* (**Table 2**). In addition, as it is frequent on birds, *A. longirostre* is popularly known as "bird tick" or "bird earring" [77].

*Amblyomma nodosum* was the second most common species collected on 12 families and 22 species of birds. This tick was recorded in co-infestation with *A. longirostre*, *A. coelebs*, *A. humerale* and *A. geayi* (**Table 2** and **Figures 2** and **4**). The birds most infested by *A. nodosum* were *Rhynchocyclus olivaceus* and *Ramphocelus carbo* with 17 and 15 nymphs, respectively. These birds inhabit the forest understory and visit the soil occasionally [67]. As Xenartha mammals (*Myrmecophaga tridactyla*,*Tamandua tetradactyla*) are the primary hosts of *A. nodosum* [17], it is believed that the low

*T. tetradactyla* may have arboreal habits [78], which helps to explain the presence of

The fauna of wild mammals in Brazil is quite diverse and more than half lives in

the Amazon biome [79]. Like birds, amphibians and reptiles, mammals play an

To date, of the total of ticks collected, 736 (70%) were larvae identified as *Amblyomma* sp. due to lack of reliable tools for larval identification, thus, indicating that the diversity of ticks on birds of the Amazon may be underestimated. Luz et al. [75], using molecular biology, identified more than 90% of the larvae collected from birds in the Atlantic Forest biome, reporting the greatest diversity of ticks on birds in a single study in Brazil and description of the new tick *Amblyomma romarioi* [14]. Therefore, the identification of all larvae by molecular biology in addition to morphological identification is extremely important to ascertain the diversity of ticks in the Amazon biome.

areas of the understory are a major source of infestation. Interestingly,

**4. Hard ticks associated with wild and domestic mammals**

*A. nodosum* also on birds of different forest strata.

(**Figure 2** and **Table 2**).

*Ecosystem and Biodiversity of Amazonia*

**Figure 4.**

**44**

**Tick species Domestic mammals Wild mammals States References**

*Cabassous unicinctus*<sup>4</sup>

*Galictis vittata*<sup>4</sup>

*vetulus*<sup>9</sup>

*, Sus*

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

*tridactyla*<sup>9</sup>

*novemcinctus*4,5,7, *Dasypus septemcinctus*4,9*, Euphractus sexcinctu*4,9*, Galactis cuja*4,9,

*Tamandua tetradactyla*<sup>4</sup>

*Thrichomys inermis*<sup>4</sup>

*Caluromys philander*<sup>5</sup>

*,*

, *Hydrochoerus*

*Cerdocyon thous*<sup>5</sup>

*marsupialis*<sup>5</sup>

*hydrochaeris*<sup>5</sup>

*gouazoubira*<sup>4</sup>

*nudicaudatus*<sup>4</sup>

*Panthera onca*<sup>5</sup>

*Pteronura brasiliensis*<sup>3</sup>

*Dasyprocta azarae*<sup>5</sup>

*Myrmecophaga tridactyla*<sup>7</sup>

*Tamandua tetradactyla*45,6

*; Didelphis albiventris*<sup>5</sup>

, *Neacomys spinosus*<sup>5</sup>

, *Pecari tajacu*<sup>7</sup>

, *Dasyprocta leporina*<sup>3</sup>

*Hydrochoerus hydrochaeris*1*,*5*,*<sup>7</sup>

, *Bradypus*

, *Bradypus variegatus*1,3,4,6,9, *Caluromys*

, *Philander*

, *Saguinus bicolor*<sup>3</sup>

,

, *Proechimys* sp<sup>3</sup>

, *Choloepus hoffmanni*<sup>7</sup>

*Tamandua tetradactyla*<sup>9</sup>

*Choloepus didactylus*3,7, *Cyclopes didactylus*<sup>3</sup>

*Tamandua tetradactyla*<sup>3</sup>

*Bradypus tridactylus*<sup>3</sup>

*Chrotopterus auritus*<sup>3</sup>

*Tamandua tetradactyla,*3,6*, Bradypus tridactylus*<sup>3</sup>

, *Dasypus novemcinctus*5,6*, Didelphis marsupialis*3,5,6*, Hydrochoerus*

*Alouatta nigerrima*<sup>6</sup>

*lituratus*<sup>3</sup>

*lanatus*<sup>3</sup>

*tridactylus*3*,*<sup>6</sup>

*marsupialis*<sup>6</sup>

*Oecomys* sp6

*didactylus*<sup>7</sup>

*opossum*<sup>3</sup>

, *Sapajus*

,*Tayassu*

,

*Didelphis marsupialis*3,5, *Hydrochoerus hydrochaeris*<sup>7</sup>

*Metachirus myosuros*<sup>5</sup>

*Bradypus tridactylus*<sup>3</sup>

*Choleopus didactylus*<sup>3</sup>

*Panthera onca*<sup>5</sup>

*macrocephalus*<sup>3</sup>

*terrestris*3,5,7

*tridactyla*5*,*<sup>9</sup>

*terrestris*<sup>5</sup>

sp7

*nasua*<sup>5</sup>

*pecari*<sup>6</sup>

sp3

*azarae*<sup>5</sup>

, *Myrmecophaga*

, *Pecari tajacu*<sup>9</sup>

, *Dasypus*

MA<sup>4</sup> , MT5 , RO<sup>7</sup> ,

[30, 32, 36, 39]

[32, 36, 42, 88–91]

[36, 38, 42]

[30–32, 38, 42, 88]

AM<sup>3</sup> [31, 36, 40]

[30, 32, 39, 54]

[30, 31, 33, 36, 39, 40, 42, 92]

, PA6 [31, 40, 42]

[30–33, 38– 40, 42, 54, 88]

TO<sup>9</sup>

AM<sup>3</sup> , MA<sup>4</sup> , MT5 , PA<sup>6</sup> ,

RO<sup>7</sup> , RR8 ,

TO<sup>9</sup>

MA<sup>4</sup> , MT5 , PA<sup>6</sup> ;

RO<sup>7</sup>

AM<sup>3</sup> , MT5 , PA<sup>6</sup> ,

RO<sup>7</sup>

,

,

, *Lycalopex*

*,*

, *Didelphis*

, *Myrmecophaga*

*, Pecari tajacu*<sup>5</sup>

, *Cuniculus paca*<sup>6</sup>

,

,*Tapirus*

,

, *Dasyprocta*

,

, *Nasua*

,

,

,*Tapirus*

, *Coendou*

, *Artibeus*

, *Didelphis*

,

, *Cyclopes*

*,*

,

,

AC<sup>1</sup> , MT5 ,

RO<sup>7</sup> , TO<sup>9</sup>

AC<sup>1</sup> , AM<sup>3</sup> , MA<sup>4</sup> , PA6 , RO<sup>7</sup> ,

TO<sup>9</sup>

AM<sup>3</sup>

AC<sup>1</sup> , AM<sup>3</sup> , MT5 , PA<sup>6</sup> ,

RO<sup>7</sup>

,

,

<sup>6</sup> *Dasyprocta*

, *Mazama*

, *Metachirus*

, *Nasua nasua*<sup>5</sup>

,*Tayassu pecari*<sup>5</sup>

*Amblyomma auricularium*

*Amblyomma cajennense* sensu stricto

*Amblyomma calcaratum*

*Amblyomma coelebs*

*Amblyomma dissimile*

*Amblyomma dubitatum*

*Amblyomma geayi*

*Amblyomma goeldii*

*Amblyomma humerale*

**47**

*Equus caballus*4*,*5*,*6*,*7*,*8*,*<sup>9</sup>

, *Canis familiaris*4*,*<sup>9</sup>

, *Bubalus*

*Equus caballus*<sup>7</sup> *Agouti paca*<sup>7</sup>

*scrofa*4*,*<sup>9</sup>

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

*bubalis*<sup>7</sup>

**Figure 5.** *Diversity of hard ticks parasitizing wild mammals from the Brazilian Amazon.*


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

**Figure 5.**

**46**

*Diversity of hard ticks parasitizing wild mammals from the Brazilian Amazon.*

*Ecosystem and Biodiversity of Amazonia*


**Tick species Domestic mammals Wild mammals States References**

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

*Equus caballus,*4*,*7*,*<sup>9</sup> *Myrmecophaga tridactyla*<sup>9</sup> MA<sup>4</sup>

*sp*3

*aguti*<sup>3</sup>

*Agouti paca*<sup>7</sup>

*gouazoubira*<sup>9</sup>

*Monodelphis glirina*<sup>6</sup>

*Monodelphis touan*<sup>6</sup>

*Bradypus tridactylus*<sup>3</sup>

*didactylus*<sup>3</sup>

*tetradactyla*<sup>3</sup>

*Marmosa murina*<sup>6</sup>

*Oligorysomys* sp.7

*demerarae*<sup>3</sup>

*opossum*<sup>6</sup>

*Mazama* p7

*Alouatta puruensis*<sup>7</sup>

*gouazoubira*5,9, *Mazama* sp.<sup>7</sup>

, *Saquinus bicolor*<sup>3</sup>

*Tapirus terrestres*7,9,*Tayassu* sp7

*azarae*<sup>5</sup>

*cuja*<sup>4</sup>

*Records of ticks in the Amazon biome, Brazil, according to tick species, hosts (domestic and wild animals),*

*Ixodes schulzei Monodelphis touan* <sup>6</sup> PA6 [92]

*Canis familiaris*4,7, *Mazama americana*3,9,

,

*Canis familiaris*1,3,4,7,9 *Bradypus tridactylus*<sup>3</sup>

*Oecomys* sp.7

*leporina*<sup>3</sup>

*pratti*<sup>1</sup>

*Ixodes lasallei Dasyprocta variegata*<sup>7</sup> RO<sup>7</sup> [16]

*Amblyomma sculptum*

*Amblyomma varium*

*Dermacentor nitens*

*Ixodes bocatorensis*

*Haemaphysalis juxtakochi*

*Rhipicephalus microplus*

*Rhipicephalus sanguineus* sensu lato

*1 Acre – AC. <sup>2</sup> Amapá – AP. <sup>3</sup> Amazonas – AM. <sup>4</sup> Maranhão – MA. <sup>5</sup> Mato Grosso – MT. <sup>6</sup> Pará – PA. <sup>7</sup> Rondônia – RO. <sup>8</sup> Roraima – RR. <sup>9</sup> Tocantins – TO.*

**Table 3.**

**49**

*states, references.*

*Bos taurus*<sup>4</sup>

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

*hircus*<sup>4</sup>

, *Canis familiaris*4,7, *Equus caballus,*4,7,8,9 *Capra*

*Ixodes amarali Hylaeamys megacephalus*<sup>6</sup>

*Ixodes luciae Didelphis marsupialis*1*,*3*,*4*,*<sup>7</sup>

, *Ovis aires*<sup>4</sup>

*Ixodes spinosus Dasyprocta* sp7

*Bos taurus*4,7,8,9*, Canis familiaris*4,7,9*, Felis catus*<sup>7</sup>

*Equus caballus,*4,7,8,9*, Capra hircus*4,7*, Ovis aires*<sup>4</sup> *Tapirus terrestris*5,7, *Hydrochoerus hydrochaeris*<sup>7</sup>

*Bradypus tridactylus*<sup>3</sup>

*Choleopus hoffmanni*3*,*<sup>7</sup>

*Choloepus* sp7 *Dasyprocta*

*, Bradypus variegatus*<sup>7</sup> *Choleopus didactylus*3*,*<sup>7</sup>

, RO<sup>7</sup> , TO<sup>9</sup>

AM<sup>3</sup> , PA6 , RO<sup>7</sup>

MA<sup>4</sup> , RO<sup>7</sup> , RR8 ,

TO<sup>9</sup>

PA6 [99]

AM<sup>3</sup> [16]

RO7, AC1 [13]

[31, 36, 38, 100]

[31, 36, 38, 39, 42]

[32, 33, 36, 38, 39, 95, 98]

[30–33, 36, 38, 39, 95, 98, 101]

*, Bradypus*

,

, *Cyclopes*

,

,

,

,

AC<sup>1</sup> , AM<sup>3</sup> , MA<sup>4</sup> , PA6 , RO<sup>7</sup>

AM<sup>3</sup> , MA<sup>4</sup> , MT5 ; RO<sup>7</sup> ,

TO<sup>9</sup>

MA<sup>4</sup> , MT5 ; RO<sup>7</sup> ,

RR8 ; TO9

AC<sup>1</sup> , AM<sup>3</sup> , MA<sup>4</sup> , RO<sup>7</sup>

*,*

, *Marmosa*

, *Philander*

, *Monodelphis* sp<sup>7</sup>

, *Orysomys* sp.7

, *Dasyprocta*

; *Dasyprocta fuliginosa*<sup>3</sup>

, *Mazama*

, *Galictis*

,

,

, *Myoprocta*

, *Dasyprocta*

,*Tamandua*

*,*

*,*

*,Tamandua tetradactyla*<sup>3</sup>

, *Mazama*

[88]

[30, 31, 33, 38, 40, 97]

[36, 38, 39, 95, 98]


**Table 3.**

**Tick species Domestic mammals Wild mammals States References** *hydrochaeris*<sup>1</sup>

*constantiae*<sup>5</sup>

*tetradactyla*3*,*<sup>6</sup>

*roberti*<sup>5</sup>

*pecari*<sup>3</sup>

*Coendou* sp<sup>3</sup>

*prehensilis*6*,*<sup>7</sup>

*nycthemera*<sup>6</sup>

*gouazoubira*<sup>4</sup>

*Agouti paca*<sup>7</sup>

*tridactyla*<sup>9</sup>

*terrestres*<sup>3</sup>

*terrestres*<sup>3</sup>

*Proechimys* sp<sup>3</sup>

*Puma concolor*<sup>5</sup>

*Tayassu pecari*<sup>7</sup>

*Dasyprocta* sp7

*Nasua nasua*<sup>7</sup>

*Cuniculus paca*<sup>5</sup>

*tetradactyla*<sup>6</sup>

*glirina*<sup>6</sup>

*Canis familiaris*<sup>7</sup> *Hydrochoerus hydrochaeris*<sup>1</sup>

*cancrivorus*<sup>7</sup>

*Dasyprocta* sp7

*Tayassu pecari*3,5,6,7, *Dasyprocta fuliginosa*<sup>3</sup>

*Mazama americana*<sup>3</sup>

*terrestris*5,7, *Panthera onca*<sup>7</sup>

*Panthera onca,*3,5,7,9,*Tapirus*

*Proechimys quadruplicatus*<sup>3</sup>

*Tayassu pecari*3,4

*acouchy*<sup>3</sup>

sp3

*Amblyomma incisum*

*Ecosystem and Biodiversity of Amazonia*

*Amblyomma latepunctatun*

*Amblyomma longirostre*

*Amblyomma naponense*

*Amblyomma nodosum*

*Amblyomma oblongoguttatum*

*Amblyomma ovale*

*Amblyomma pacae*

*Amblyomma parkeri*

*Amblyomma romitii*

*Amblyomma rotundatum*

*Amblyomma scalpturatum*

**48**

*Canis familiaris*4,6,9, *Sus*

*Canis familiaris*3,4,9, *Sus*

*Canis familiaris*<sup>7</sup>

*scrofa*<sup>7</sup>

*, Sus*

*scrofa*<sup>7</sup>

*scrofa*<sup>7</sup>

*Metachirus myosuros*<sup>5</sup>

*,Tamandua*

*Philander opossum*<sup>3</sup>

*Tapirus terrestris*3*,*6*,*<sup>7</sup>

*Didelphis marsupialis*<sup>3</sup>

*Dasyprocta fuliginosa*<sup>3</sup>

, *Hydrochoerus hydrochaeris*<sup>1</sup>

, *Marmosa*

, *Marmosa* sp<sup>3</sup>

, *Dasyprocta fuliginosa*<sup>3</sup>

, *Coendou*

, *Coendou*

, *Mazama*

,*Tapirus terrestris*<sup>7</sup>

*Tamandua tetradactyla*4,5,6,7,9; *Bradypus variegatus*<sup>9</sup>

*Myrmecophaga tridactyla*<sup>9</sup>

, *Myoprocta*

,

*, Cuniculus paca*<sup>9</sup>

,*Tayassu tacaju*<sup>7</sup>

, *Myrmecophaga*

,

, *Leopardus pardalis*<sup>3</sup>

,*Tapirus*

, *Procyon*

,*Tayassu tacaju*<sup>7</sup>

*, Agouti paca*<sup>7</sup>

*, Eira barbara*<sup>7</sup>

, *Cerdocyon thous*<sup>9</sup>

,*Tamandua*

, *Monodelphis*

*, Didelphis marsupialis*<sup>7</sup>

*Hydrochoerus hydrochaeris*5,6,7 MT5

*, Dasyprocta*

, *Pecari tajacu*5,7,

*, Didelphis*

*Hydrochoerus hydrochaeris<sup>i</sup>*

*Xenodon merremii*<sup>9</sup>

*Agouti paca*<sup>7</sup>

*marsupialis*<sup>5</sup>

*fuliginosa*<sup>3</sup>

*, Pauxi tuberosa*<sup>5</sup>

,*Tapirus*

*,*

,

, *Proechimys*

*Tapirus terrestris*<sup>7</sup> RO<sup>7</sup> [38, 93]

,

AM<sup>3</sup> ; PA6 ; RO<sup>7</sup>

AM<sup>3</sup> , PA6 , RO<sup>7</sup>

AC<sup>1</sup> , AM<sup>3</sup> , MA<sup>4</sup> , RO<sup>7</sup>

MA<sup>4</sup> , MT5 , PA<sup>6</sup> ,

RO<sup>7</sup> , TO<sup>9</sup>

AM<sup>3</sup> , MA<sup>4</sup> , MT5 , PA<sup>6</sup> ,

RO<sup>7</sup> , TO<sup>9</sup>

AM<sup>3</sup> , MA<sup>4</sup> , MT5 , RO<sup>7</sup> ,

TO<sup>9</sup> ,

AC<sup>1</sup> , MT5 ,

PA6 , RO<sup>7</sup>

RO<sup>7</sup>

AM3, MT5 , RO<sup>7</sup>

, PA<sup>6</sup> ,

RO7, TO9 [30–33, 36, 38, 39, 41, 42, 54]

,

,

*,*

,

,

*,*

*Didelphis marsupialis*<sup>5</sup> MT5 [32]

*,*

,

*,*

*,*

[31, 38, 93]

[30, 31, 33, 40, 42]

[31, 36, 38, 54]

[30, 32, 38– 40, 42]

[31, 32, 38, 39, 42, 91, 94, 95]

[30–32, 38, 39, 42, 91, 94]

[38, 42, 54, 92]

[32, 41, 96]

[30–32, 38, 88]

,*Tayassu*

, *Galictis*

,

*Records of ticks in the Amazon biome, Brazil, according to tick species, hosts (domestic and wild animals), states, references.*

rodents (e.g., Cricetidae) in the larvae and nymph stages, with adults feeding mostly on marsupials (e.g., *Didelphis* spp., *Monodelphis* spp.) [17, 83], and agoutis (e.g., *Dasyprocta*) [13, 16]. Most species were found parasitizing *Monodelphis glirina* (*I. amarali*, *I. schulzei*, *I. luciae*) and *Monodelphis touan* (*I. amarali*, *I. schulzei*)*,* with some reports of *I. amarali* on the rodent *Hylaeamys megacephalus* [30, 38, 92, 99, 100]. Also on rodents the species *I. spinosus, I. lasallei* and *I. bocatorensis* have been recorded [30, 31, 38], although these previous records referred to them erroneously as *Ixodes fuscipes*, which according to more recent data, does not occur in the Amazon biome (**Figure 5** and **Table 3**) [13, 16].

of Santa Isabel do Rio Negro [31, 38]. *H. juxtakochi* has been found infected with the bacterium *Rickettsia rhipicephali* in the Amazon region, more precisely in the municipalities of Monte Negro and Confresa, states of Rondônia and Mato Grosso, respectively [42, 93]. Human parasitism by *R. microplus* was expected, since this species can parasitize humans when it comes into direct contact with domestic cattle, as reported by [38]. Therefore, its importance is much more economical,

Despite sporadic records of *R. sanguineus* s.l. on humans in Brazil, this association in the Brazilian Amazon requires confirmation. Records of approximately 4,020 *R. sanguineus s.l.* (larva, nymph, and adult), supposedly collected on humans, in the state of Pará by Serra-Freire et al. [103] do not corroborate the common findings in country. The bioecology of this endophilic and introduced tick is well studied. In other countries, *R. sanguineus* s.l. is vector of some zoonotic agents for humans (*Rickettsia conorii*, *Rickettsia massiliae* and *R. rickettsii*) [83]. Adults of *A. cajennense* s.s. have been reported on humans in the municipality of Monte Alegre (Pará state) [90]. Reports of *Amblyomma cajennense* sensu lato parasitizing humans published by Martins et al. [90] in the municipalities of Sinop and Tucuruí in the states of Mato Grosso and Pará, is possibly *A. cajennense* s.s. taking into account the area of occurrence of this species. This tick is aggressive to humans. In the Amazon biome the *R. bellii* bacterium was detected in this species in Mato Grosso state and *R. amblyommatis* in Mato Grosso, Maranhão and Rondônia state [32, 42, 53, 83,

Nymphs and adult of *A. coelebs* were found on humans in Rondônia state [38]. An adult of this tick was found on human in Roraima state, municipality of Caroebe [106]. The nymph and adult stages of *A. colelebs* were also found on humans in Amazonas state, in the municipalities of Coari, Fonte Boa, Presidente Figueiredo and Santa Isabel do Rio Negro [31, 107]. There are reports of infection by *R. amblyommatis* in *A. coelebs* in the Amazon region in the states of Rondônia and Mato Grosso, respectively [32, 53, 88]. Adult of *A. dissimile* was recorded on humans in the Maicurú River/Amazon biome, located in Pará state [106]. Adults of the species *A. latepunctatum* have been reported on humans in Coari and Santa Isabel do Rio

Adults of the tick *A. naponense* (published as *Amblyomma mantiquirense*) were found on humans in the state of Pará (municipality not reported) [108]. Human parasitism also by adults was later reported on the Maicurú River, located in this same state [106]. However, nymphs have been found on humans in the states of Rondônia (municipality not specified precisely) and Amazonas, in the municipalities of Coari and Santa Isabel do Rio Negro [31, 38]. The bacteria *R. bellii* and *Rickettsia* sp. strain PA were identified in this species in the Amazon region of the municipalities of Santarém and Rurópolis, in the state of Pará [42]. *Rickettsia* sp. strain Tapirapé was found in this tick in the municipalities of Confresa and Rio

Adults of the species *A. oblongoguttatum* were recorded parasitizing humans in the state of Pará, in the Maicurú River and in the municipality of Uruará [106, 108, 109]. Human parasitism by nymphs and adults of this tick was recorded in the state

Negro, both municipalities located in the state of Amazonas [31].

Branco, in the states of Mato Grosso and Acre, respectively [42, 54].

of Rondônia (municipality not specified with precision) [38]. However, the nymphal stage of this species was later recorded on humans in the municipality of Monte Negro in the same state [106]. Human parasitism by adult of this tick was also recorded in the municipality of Caroebe, state of Roraima [106]. Nymphs and adults of this species have been recorded parasitizing humans in three municipalities (Coari, Manacapuru and Santa Isabel do Rio Negro), all located in the state of Amazonas [31, 106, 107]. On the other hand, the life cycle of this tick has already

been studied under laboratory conditions [110]. The bacteria *R. bellii* and

causing severe losses to livestock in the country than for public health.

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

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

88, 90, 91, 104, 105].

**51**

In general, the greatest diversity of ticks was recorded on *T. tetradactyla* with 10 species, followed by the rodent *H. hydrochaeris* with nine species. These records indicates the importance of these hosts for maintaining local tick diversity, in addition to act as dispersers of these ectoparasites. Additional hosts have also been shown to be important in maintaining diversity of tick in the Amazon: *Tapirus terrestris*,*Tayassu pecari*, *Pecari tajacu*, *Agouti paca*, *Dasyprocta fuliginosa*, *Bradypus tridactylus*, *Bradypus variegatus*, *Panthera onca*, *Nasua nasua*, *Choleopus didactylus*, *Choloepus hoffmanni*, *Cyclopes didactylus*, *Dasyprocta azarae*, *Dasyprocta leporina*, *Philander opossum*, *M. glirina*, *M. touan*, *Mazama gouazoubira*, *Mazama americana* and *M. tridactyla* (**Table 3**), because they are parasitized by more than one species of ticks. There is also a record of *A. cajennense s.s.* on *Pteronura brasiliensis* a semiaquatic animal [89].

Interestingly, three species of ticks commonly found on domestic animals have also been found parasitizing wild animals: *Rhipicephalus sanguineus* sensu lato, *R. microplus* and *D. nitens*. Although occasional, the encounter of these species of ticks on wild animals is possible, especially when wild animals coexist with domestic animals infested by ticks (e.g., pastures, corrals, houses). A total of 11 species of ticks have been recorded parasitizing domestic animals in Amazon (**Table 3**).

#### **5. Hard ticks associate with humans**

Spotted fevers caused by *R. rickettsii* and *Rickettsia parkeri* are currently the only confirmed tick-borne disease affecting humans in the country, and *A. sculptum*, *A. aureolatum*, *A. ovale* are the main vectors. The disease is severe and highly lethal when caused by the bacterium *R. rickettsii* (vectors *A. sculptum* and *A. aureolatum*) and moderate, non-lethal, when caused by *R. parkeri*, vectored mainly by *A. ovale* [10]. Only *A. sculptum* and *A. ovale* have been recorded in the Brazilian Amazon, the first in rare reports and the second frequently recorded and populations established in this biome. In fact, considering only the tick adult stage, *A. ovale* has been reported as the most common human-biting tick in the Amazon [38].

In the last century, papers about ticks on human beings have been published in the Brazilian Amazon, however these studies are fragmented and scarce. At least 14 species of hard ticks have already been found and documented parasitizing humans within the limits of the Brazilian Amazon. Of these, the vast majority (11 species) belong to the genus *Amblyomma*: *A. cajennense* s.s., *A. coelebs*, *A. dissimile*, *A. latepunctatum*, *A. naponense, A. oblongoguttatum*, *A. ovale*, *A. romitii*, *A. rotundatum*, *A. scalpturatum* and *A. sculptum*. The other three species belong to the genus *Rhipicephalus,* including *R. microplus* and *R. sanguineus* s.l.*,* and the genus *Haemaphysalis* with only *H. juxtakochi*. The public health importance of human–tick associations in the Amazon biome is unclear and further research are needed to clarify the issue.

Nymphs of *H. juxtakochi* have been reported on humans in the states of Rondônia (municipality not precisely indicated) and Amazonas, in the municipality

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

of Santa Isabel do Rio Negro [31, 38]. *H. juxtakochi* has been found infected with the bacterium *Rickettsia rhipicephali* in the Amazon region, more precisely in the municipalities of Monte Negro and Confresa, states of Rondônia and Mato Grosso, respectively [42, 93]. Human parasitism by *R. microplus* was expected, since this species can parasitize humans when it comes into direct contact with domestic cattle, as reported by [38]. Therefore, its importance is much more economical, causing severe losses to livestock in the country than for public health.

Despite sporadic records of *R. sanguineus* s.l. on humans in Brazil, this association in the Brazilian Amazon requires confirmation. Records of approximately 4,020 *R. sanguineus s.l.* (larva, nymph, and adult), supposedly collected on humans, in the state of Pará by Serra-Freire et al. [103] do not corroborate the common findings in country. The bioecology of this endophilic and introduced tick is well studied. In other countries, *R. sanguineus* s.l. is vector of some zoonotic agents for humans (*Rickettsia conorii*, *Rickettsia massiliae* and *R. rickettsii*) [83]. Adults of *A. cajennense* s.s. have been reported on humans in the municipality of Monte Alegre (Pará state) [90]. Reports of *Amblyomma cajennense* sensu lato parasitizing humans published by Martins et al. [90] in the municipalities of Sinop and Tucuruí in the states of Mato Grosso and Pará, is possibly *A. cajennense* s.s. taking into account the area of occurrence of this species. This tick is aggressive to humans. In the Amazon biome the *R. bellii* bacterium was detected in this species in Mato Grosso state and *R. amblyommatis* in Mato Grosso, Maranhão and Rondônia state [32, 42, 53, 83, 88, 90, 91, 104, 105].

Nymphs and adult of *A. coelebs* were found on humans in Rondônia state [38]. An adult of this tick was found on human in Roraima state, municipality of Caroebe [106]. The nymph and adult stages of *A. colelebs* were also found on humans in Amazonas state, in the municipalities of Coari, Fonte Boa, Presidente Figueiredo and Santa Isabel do Rio Negro [31, 107]. There are reports of infection by *R. amblyommatis* in *A. coelebs* in the Amazon region in the states of Rondônia and Mato Grosso, respectively [32, 53, 88]. Adult of *A. dissimile* was recorded on humans in the Maicurú River/Amazon biome, located in Pará state [106]. Adults of the species *A. latepunctatum* have been reported on humans in Coari and Santa Isabel do Rio Negro, both municipalities located in the state of Amazonas [31].

Adults of the tick *A. naponense* (published as *Amblyomma mantiquirense*) were found on humans in the state of Pará (municipality not reported) [108]. Human parasitism also by adults was later reported on the Maicurú River, located in this same state [106]. However, nymphs have been found on humans in the states of Rondônia (municipality not specified precisely) and Amazonas, in the municipalities of Coari and Santa Isabel do Rio Negro [31, 38]. The bacteria *R. bellii* and *Rickettsia* sp. strain PA were identified in this species in the Amazon region of the municipalities of Santarém and Rurópolis, in the state of Pará [42]. *Rickettsia* sp. strain Tapirapé was found in this tick in the municipalities of Confresa and Rio Branco, in the states of Mato Grosso and Acre, respectively [42, 54].

Adults of the species *A. oblongoguttatum* were recorded parasitizing humans in the state of Pará, in the Maicurú River and in the municipality of Uruará [106, 108, 109]. Human parasitism by nymphs and adults of this tick was recorded in the state of Rondônia (municipality not specified with precision) [38]. However, the nymphal stage of this species was later recorded on humans in the municipality of Monte Negro in the same state [106]. Human parasitism by adult of this tick was also recorded in the municipality of Caroebe, state of Roraima [106]. Nymphs and adults of this species have been recorded parasitizing humans in three municipalities (Coari, Manacapuru and Santa Isabel do Rio Negro), all located in the state of Amazonas [31, 106, 107]. On the other hand, the life cycle of this tick has already been studied under laboratory conditions [110]. The bacteria *R. bellii* and

rodents (e.g., Cricetidae) in the larvae and nymph stages, with adults feeding mostly on marsupials (e.g., *Didelphis* spp., *Monodelphis* spp.) [17, 83], and agoutis (e.g., *Dasyprocta*) [13, 16]. Most species were found parasitizing *Monodelphis glirina* (*I. amarali*, *I. schulzei*, *I. luciae*) and *Monodelphis touan* (*I. amarali*, *I. schulzei*)*,* with some reports of *I. amarali* on the rodent *Hylaeamys megacephalus* [30, 38, 92, 99, 100]. Also on rodents the species *I. spinosus, I. lasallei* and *I. bocatorensis* have been recorded [30, 31, 38], although these previous records referred to them erroneously as *Ixodes fuscipes*, which according to more recent data, does not occur in the

In general, the greatest diversity of ticks was recorded on *T. tetradactyla* with 10 species, followed by the rodent *H. hydrochaeris* with nine species. These records indicates the importance of these hosts for maintaining local tick diversity, in addition to act as dispersers of these ectoparasites. Additional hosts have also been shown to be important in maintaining diversity of tick in the Amazon: *Tapirus terrestris*,*Tayassu pecari*, *Pecari tajacu*, *Agouti paca*, *Dasyprocta fuliginosa*, *Bradypus tridactylus*, *Bradypus variegatus*, *Panthera onca*, *Nasua nasua*, *Choleopus didactylus*, *Choloepus hoffmanni*, *Cyclopes didactylus*, *Dasyprocta azarae*, *Dasyprocta leporina*, *Philander opossum*, *M. glirina*, *M. touan*, *Mazama gouazoubira*, *Mazama americana* and *M. tridactyla* (**Table 3**), because they are parasitized by more than one species of ticks. There is also a record of *A. cajennense s.s.* on *Pteronura brasiliensis* a semi-

Interestingly, three species of ticks commonly found on domestic animals have

Spotted fevers caused by *R. rickettsii* and *Rickettsia parkeri* are currently the only confirmed tick-borne disease affecting humans in the country, and *A. sculptum*, *A. aureolatum*, *A. ovale* are the main vectors. The disease is severe and highly lethal when caused by the bacterium *R. rickettsii* (vectors *A. sculptum* and *A. aureolatum*) and moderate, non-lethal, when caused by *R. parkeri*, vectored mainly by *A. ovale* [10]. Only *A. sculptum* and *A. ovale* have been recorded in the Brazilian Amazon, the first in rare reports and the second frequently recorded and populations established in this biome. In fact, considering only the tick adult stage, *A. ovale* has

In the last century, papers about ticks on human beings have been published in the Brazilian Amazon, however these studies are fragmented and scarce. At least 14 species of hard ticks have already been found and documented parasitizing humans within the limits of the Brazilian Amazon. Of these, the vast majority (11 species) belong to the genus *Amblyomma*: *A. cajennense* s.s., *A. coelebs*, *A. dissimile*, *A. latepunctatum*, *A. naponense, A. oblongoguttatum*, *A. ovale*, *A. romitii*, *A. rotundatum*, *A. scalpturatum* and *A. sculptum*. The other three species belong to the genus *Rhipicephalus,* including *R. microplus* and *R. sanguineus* s.l.*,* and the genus

*Haemaphysalis* with only *H. juxtakochi*. The public health importance of human–tick associations in the Amazon biome is unclear and further research are needed to

Nymphs of *H. juxtakochi* have been reported on humans in the states of Rondônia (municipality not precisely indicated) and Amazonas, in the municipality

been reported as the most common human-biting tick in the Amazon [38].

also been found parasitizing wild animals: *Rhipicephalus sanguineus* sensu lato, *R. microplus* and *D. nitens*. Although occasional, the encounter of these species of ticks on wild animals is possible, especially when wild animals coexist with domestic animals infested by ticks (e.g., pastures, corrals, houses). A total of 11 species of ticks have been recorded parasitizing domestic animals in Amazon (**Table 3**).

Amazon biome (**Figure 5** and **Table 3**) [13, 16].

*Ecosystem and Biodiversity of Amazonia*

**5. Hard ticks associate with humans**

aquatic animal [89].

clarify the issue.

**50**

*R. amblyommatis* were detected in this species in the Amazon in the municipalities of Governador Jorge Teixeira and Pimenta Bueno, in the state of Rondônia [53, 105].

and adults in the rainy period (October to March) [119]. Similar to the pattern

fication of soft ticks into five genera, namely: *Antricola, Argas, Nothoaspis,*

over massive rock formations where bats shelter (**Table 4**).

The fauna of argasid ticks in Brazil is currently composed by 24 species [15]. Ticks of this family parasitize terrestrial vertebrates including amphibians in this country [25, 26]. With the exception of larvae from an undetermined *Ornithodoros* sp. collected on *Potus flavus* (Procionidae) [84], reports of soft ticks parasitizing mammals in the Brazilian Amazon are few and almost restricted to larvae collected on bats. Adults and nymphs have been collected either inside bat-inhabited caves or

The first record of a soft tick in the Brazilian Amazon was published by North American entomologists Robert A. Cooley and Glen M. Kohls back in 1941. They received a tick collected inside a bat-inhabited three hole at Marajó Island (Pará state) and identified it as female of *Ornithodoros hasei* (mentioned as *Ornithodoros dunni*) [126]. Nowadays, we know that *O. hasei* is a wide spread species in Brazil and that three-roosting bats *Artibeus planirostris* and *Noctilio* spp*.* could act as main

*Tadarida laticaudata* were the first bats reported to be parasitized by soft ticks in the Brazilian Amazon [133]. This report referred to *Ornithodoros setosus,* which was recently reclassified as *Nothoaspis setosus* [124]. In 1972, the bats *Noctilio labialis* and *T. laticaudata* were reported to be parasitized by *Ornithodoros stageri* in the Brazilian Amazon [129], which was recently confirmed [11]. A report in the Brazilian Amazon is a remarkable fact for *O. stageri*, since it also has distribution in Southern

Bat inhabited caves constitute excellent niches to find argasid ticks. In particular, special, large colonies of insectivorous bats dwelling inside small chambers create high temperature conditions (28-40°C) where hundreds of *Antricola*, *Nothoaspis* and *Ornithodoros* ticks might occur [135]. Between 2004 to 2010 collections of ticks performed in hot caves from Porto Velho (Rondônia) fostered the description of two novel species (e.g., *Nothoaspis amazoniensis* and *Ornithodoros rondoniensis*) [122, 123]; the redescription of larvae and description of postlarval stages of *Ornithodoros marinkellei* [127], and the expansion of geographical distribution of *Antricola delacruzi* and *Antricola guglielmonei* into Brazilian Amazon [122]. At least for two species, *O. marinkellei* and *O. rondoniensis*, further collections performed in caves from Pará state underpined a larger distribution along the

Most amazing feature of soft ticks inhabiting hot caves, is that adaptation to this particular milieu seems to have modified their morphology and biology drastically. For instance, evidence showing that adults of *Antricola* ticks lack the capacity to

hosts [87, 130–132]. With the exception of recent collection of larvae on *A. planirostris* [87], knowledge on the distribution of *O. hasei* along the Brazilian

Taxonomy of Argasidae is currently questionable and relies on five schemes that divide the family in up to ten genera [120]. Achieving a consensus between soft tick taxonomists depends now chiefly on the molecular characterization of early collected type specimens for some genera. For instance, elucidating the status of pivotal taxa such as *Alectorobius* is mandatory if we are to understand the systematics of the Argasidae, particularly in the American Continent. Considering a practical approach, in this chapter we adopt but not necessarily endorse the classi-

observed for *A. sculptum* in areas outside the Amazon biome [116].

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

**6. Soft ticks in the Brazilian Amazon**

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

*Ornithodoros*, and *Otobius* [121].

Amazon is still poorly vague.

Amazon ecosystems [128].

**53**

United States and Mexico [126, 134].

Adults of the tick *A. ovale* have been reported to parasitize humans in several areas of the state of Rondônia [38]. As previously reported in this chapter, adults of *A. ovale* parasitize mainly domestic and wild carnivores, while immature (larva and nymph) parasitize rodents of the families Cricetidae and Echimyidae, with sporadic reports on wild birds [17, 75, 83]. This preference for such host groups was observed in the laboratory [111]. This tick is a vector of the protozoan *Hepatozoon canis* and the bacterium *R. parkeri*, agents of importance in veterinary and human medicine, respectively [83]. In the Amazon region, the bacterium *R. bellii* was identified in this species in the municipalities of Governador Jorge Teixeira and Monte Negro, in the state of Rondônia; and in the municipality of Cururupu, in the state of Maranhão [53, 91].

The adult stage of the species *A. romitii* (published as *Amblyomma tasquei*) was found parasitizing humans in the north of the state of Pará (municipality not reported) [108]. In this same state, the larval and adult stages of this tick were found on humans in the municipality of Rurópolis [96, 112]. The life cycle of this tick was studied under laboratory conditions [113].

Parasitism by adults of *A. rotundatum* was recorded in the municipality of Belém in the state of Pará on a human who worked in frequent contact with reptiles in the Zoobotanical Park of the Museu Paraense Emílio Goeldi [114].

Adults of the *A. scalpturatum* have been reported to parasitize humans in the state of Pará (municipality not reported) [108], and in the Maicurú River located in this state [93, 106]. Human parasitism by the nymphal and adult stages of this species has been reported in the states of Mato Grosso (municipality of Jauru) and Rondônia (municipality not precisely indicated), however in this latter state it was not specified whether these adult ticks were fixed or walking on humans [38, 93]. The nymphal stage was also collected from a human who was probably infested with this tick in the municipality of Porto Velho in the state of Rondônia [115]. The nymph and adult stages of this species were also found parasitizing humans in the state of Amazonas, in the municipalities of Coari, Fonte Boa, Jutaí, Presidente Figueiredo and Santa Isabel Rio Negro [31, 107]. In the Amazon biome, two species of rickettsiae (*R. bellii* and *R. amblyommatis*) were detected in *A. scalpturatum,* in the municipalities of Governador Jorge Teixeira and Sinop, states of Rondônia and Mato Grosso, respectively [53, 88].

Human parasitism by adults of *A. sculptum* was recorded in the municipality of Jauru in the state of Mato Grosso [90]. The Jauru territory comprises 90% of the Amazon biome and 10% of the Cerrado biome, so this human record in this region is in accordance with the transition area of the occurrence of this species in sympatry with *A. cajennense* s. s. [90]. The bioecology of this native tick has been extensively studied due to its aggressive parasitism towards domestic animals and humans in the Brazilian territory. In the geographical area in which this species occurs, it completes an annual generation, with larvae occurring mostly during autumn, nymphs during winter, and adults during spring and summer, with larval behavioral diapause being the main regulating factor of its life cycle [116, 117]. Besides being considered a pest for domestic animals (dogs, cattle and horses), the species can transmit to humans the Brazilian Spotted Fever caused by the bacterium *R. rickettsii*, being this the most important zoonotic disease transmitted by ticks within the limits of the National territory. Additionally, *A. sculptum* has also been identified with the bacterium *R. amblyommatis* in the municipality of Pium, state of Tocantins [118].

There is a trend of seasonal behavior for some species of *Amblyomma* in the Amazon biome, with immatures predominating in the dry period (June to October), and adults in the rainy period (October to March) [119]. Similar to the pattern observed for *A. sculptum* in areas outside the Amazon biome [116].
