# Coxiella burnetii Infection in Livestock, Pets, Wildlife, and Ticks in Latin America and the Caribbean: a Comprehensive Review of the Literature

Loïc Epelboin, Mateus de Souza Ribeiro Mioni, Aurelie Couesnon, Mona Saout, Edith Guilloton, Salma Omar, Vincent Pommier de Santi, Bernard Davoust, Jean Lou Marié, Anne Lavergne, et al.

<figure><img src="image_1.png" /></figure>

## ► To cite this version:

Loïc Epelboin, Mateus de Souza Ribeiro Mioni, Aurelie Couesnon, Mona Saout, Edith Guilloton, et al.. Coxiella burnetii Infection in Livestock, Pets, Wildlife, and Ticks in Latin America and the Caribbean: a Comprehensive Review of the Literature. Current tropical medicine reports, 2023, 10 (3), pp.94-137. 10.1007/s40475-023-00288-7. hal-04275614

HAL Id: hal-04275614

https://hal.science/hal-04275614

Submitted on 17 Apr 2024

HAL is a multi-disciplinary open access
archive for the deposit and dissemination of sci-
entific research documents, whether they are pub-
lished or not. The documents may come from
teaching and research institutions in France or
abroad, or from public or private research centers.

L'archive ouverte pluridisciplinaire HAL, est
destinée au dépôt et à la diffusion de documents
scientifiques de niveau recherche, publiés ou non,
émanant des établissements d'enseignement et de
recherche français ou étrangers, des laboratoires
publics ou privés.

<figure><img src="image_2.png" /></figure>

Distributed under a Creative Commons Attribution 4.0 International License
# Coxiella burnetii Infection in Livestock, Pets, Wildlife, and Ticks in Latin America and the Caribbean: a Comprehensive Review of the Literature

Loïc Epelboin, Mateus de Souza Ribeiro Mioni, Aurelie Couesnon, Mona Saout, Edith Guilloton, Salma Omar, Vincent Pommier de Santi, Bernard Davoust, Jean Lou Marié, Anne Lavergne, et al.

<figure><img src="image_1.png" /></figure>

## ► To cite this version:

Loïc Epelboin, Mateus de Souza Ribeiro Mioni, Aurelie Couesnon, Mona Saout, Edith Guilloton, et al.. Coxiella burnetii Infection in Livestock, Pets, Wildlife, and Ticks in Latin America and the Caribbean: a Comprehensive Review of the Literature. Current tropical medicine reports, 2023, 10 (3), pp.94-137. 10.1007/s40475-023-00288-7. hal-04275614

HAL Id: hal-04275614

https://hal.science/hal-04275614v1

Submitted on 17 Apr 2024

HAL is a multi-disciplinary open access
archive for the deposit and dissemination of sci-
entific research documents, whether they are pub-
lished or not. The documents may come from
teaching and research institutions in France or
abroad, or from public or private research centers.

L'archive ouverte pluridisciplinaire HAL, est
destinée au dépôt et à la diffusion de documents
scientifiques de niveau recherche, publiés ou non,
émanant des établissements d'enseignement et de
recherche français ou étrangers, des laboratoires
publics ou privés.

<figure><img src="image_2.png" /></figure>

Distributed under a Creative Commons Attribution 4.0 International License
# *Coxiella burnetii* Infection in Livestock, Pets, Wildlife, and Ticks in Latin America and the Caribbean: a Comprehensive Review of the Literature

Loïc Epelboin¹,² • Mateus De Souza Ribeiro Mioni³ • Aurelie Couesnon⁴ • Mona Saout⁵ • Edith Guilloton² •

Salma Omar² • Vincent Pommier De Santi⁶ • Bernard Davoust⁷,⁸ • Jean Lou Marié⁷,⁹ • Anne Lavergne¹⁰ •

Damien Donato¹⁰ • Alejandro Guterres¹¹ • Sebastien Rabier² • Justin Destoop¹² • Felix Djossou¹ •

Xavier Baudrimont¹³ • Antoine Roch¹³ • Gabriel Leonardo Cicuttin¹⁴ • Tatiana Rozental¹¹,¹⁵ • Mathieu Nacher² •

Javier Millán¹⁶,¹⁷,¹⁸ • Elba R. Sampaio De Lemos¹¹ • Jorlan Fernandes¹¹ • Olivier Duron¹⁹ • Benoit De Thoisy¹⁰,²⁰ •

Elodie Rousset⁴

Accepted: 17 April 2023 / Published online: 15 June 2023
© The Author(s) 2023

## Abstract

**Purpose of the Review** Q fever, a bacterial zoonosis caused by *Coxiella burnetii*, is reported very heterogeneously in humans in Latin America. The objective of this study was to review the data on *Coxiella burnetii* Infection in animals in Latin America and the Caribbean.

**Recent Findings** A comprehensive literature review was carried out in the 47 countries and territories of Latin America on various search engines and grouped into four groups: livestock, pets, wildlife, and ticks.

**Summary** Thus, 113 studies were selected between 1950 and 2022. Among the 47 countries, only 25 (53%) had at least one publication related to *C. burnetii* infection in animals. The most productive country was Brazil (N=51), followed by French Guiana (N=21), and Colombia (N=16). Studies in livestock from 20 countries have shown widely varying country-to-country rates of seroprevalence, ranging from 0 to 67%. Some studies from seven countries, especially French Guiana and Brazil, found antibodies and sometimes positive PCR in dogs and cats, generally in the context of investigations around human clustered cases. Knowledge remained fragmented about infection in wildlife from only five countries (Chile, Colombia, Brazil, French Guiana, and Uruguay). *C. burnetii* infection was identified by PCR in Chiroptera (7 species), Rodentia (6 species), Suina (2 species), Xenartha (1 species), Cingulata (1 species), and Perissodactyla (1 species). Studies on *Coxiella* sp. in ticks have been performed in 11 countries, mostly in Brazil, and mainly found *Coxiella*-like endosymbionts. Thus, data on *C. burnetii* infection in animals are sparse and incomplete in Latin America and the Caribbean, and more research is warranted.

**Keywords** *Coxiella burnetii* • Latin America • Caribbean • Animal health • Zoonosis • Wildlife • Ticks • Cattle

## Introduction

Q fever is a zoonosis caused by the strict intracellular bacterium *Coxiella burnetii*, which is maintained in the environment by animals, especially mammals [1–3]. Humans generally become infected through air-borne transmission of the bacterium from the environment contaminated by animal reservoirs, as a spore-like form allows *C. burnetii* to resist

different environmental conditions and disinfectants [4]. This zoonosis is found worldwide, but there is considerable uncertainty about the incidence in different regions of the world and the relative importance of the risk factors of human infection. Potential reservoirs of the pathogen are numerous in wild and domestic species [5–8]. Nevertheless, descriptive studies mainly targeted -cattle, sheep, and goats - which represent the main source of human infection in Europe, while cattle appear to be involved as major reservoirs in Canada and Japan [1, 9, 10]. Other domestic mammals, such as cats and dogs, may also play a role in the transmission to humans, for instance in Canada and Australia [11–17]. Although less frequent, birds, marine mammals, or game may be sometimes implicated in the occurrence of sporadic or clustered human cases [17–20].

This article is part of the Topical Collection on *Emerging Tropical Diseases*.

Extended author information available on the last page of the article
# *Coxiella burnetii* Infection in Livestock, Pets, Wildlife, and Ticks in Latin America and the Caribbean: a Comprehensive Review of the Literature

Loïc Epelboin¹,² • Mateus De Souza Ribeiro Mioni³ • Aurelie Couesnon⁴ • Mona Saout⁵ • Edith Guilloton² •

Salma Omar² • Vincent Pommier De Santi⁶ • Bernard Davoust⁷,⁸ • Jean Lou Marié⁷,⁹ • Anne Lavergne¹⁰ •

Damien Donato¹⁰ • Alejandro Guterres¹¹ • Sebastien Rabier² • Justin Destoop¹² • Felix Djossou¹ •

Xavier Baudrimont¹³ • Antoine Roch¹³ • Gabriel Leonardo Cicuttin¹⁴ • Tatiana Rozental¹¹,¹⁵ • Mathieu Nacher² •

Javier Millán¹⁶,¹⁷,¹⁸ • Elba R. Sampaio De Lemos¹¹ • Jorlan Fernandes¹¹ • Olivier Duron¹⁹ • Benoit De Thoisy¹⁰,²⁰ •

Elodie Rousset⁴

Accepted: 17 April 2023 / Published online: 15 June 2023

© The Author(s) 2023

## Abstract

**Purpose of the Review** Q fever, a bacterial zoonosis caused by *Coxiella burnetii*, is reported very heterogeneously in humans in Latin America. The objective of this study was to review the data on *Coxiella burnetii* Infection in animals in Latin America and the Caribbean.

**Recent Findings** A comprehensive literature review was carried out in the 47 countries and territories of Latin America on various search engines and grouped into four groups: livestock, pets, wildlife, and ticks.

**Summary** Thus, 113 studies were selected between 1950 and 2022. Among the 47 countries, only 25 (53%) had at least one publication related to *C. burnetii* infection in animals. The most productive country was Brazil (N=51), followed by French Guiana (N=21), and Colombia (N=16). Studies in livestock from 20 countries have shown widely varying country-to-country rates of seroprevalence, ranging from 0 to 67%. Some studies from seven countries, especially French Guiana and Brazil, found antibodies and sometimes positive PCR in dogs and cats, generally in the context of investigations around human clustered cases. Knowledge remained fragmented about infection in wildlife from only five countries (Chile, Colombia, Brazil, French Guiana, and Uruguay). *C. burnetii* infection was identified by PCR in Chiroptera (7 species), Rodentia (6 species), Suina (2 species), Xenartha (1 species), Cingulata (1 species), and Perissodactyla (1 species). Studies on *Coxiella* sp. in ticks have been performed in 11 countries, mostly in Brazil, and mainly found *Coxiella*-like endosymbionts. Thus, data on *C. burnetii* infection in animals are sparse and incomplete in Latin America and the Caribbean, and more research is warranted.

**Keywords** *Coxiella burnetii* • Latin America • Caribbean • Animal health • Zoonosis • Wildlife • Ticks • Cattle

## Introduction

Q fever is a zoonosis caused by the strict intracellular bacterium *Coxiella burnetii*, which is maintained in the environment by animals, especially mammals [1–3]. Humans generally become infected through air-borne transmission of the bacterium from the environment contaminated by animal reservoirs, as a spore-like form allows *C. burnetii* to resist

different environmental conditions and disinfectants [4]. This zoonosis is found worldwide, but there is considerable uncertainty about the incidence in different regions of the world and the relative importance of the risk factors of human infection. Potential reservoirs of the pathogen are numerous in wild and domestic species [5–8]. Nevertheless, descriptive studies mainly targeted -cattle, sheep, and goats - which represent the main source of human infection in Europe, while cattle appear to be involved as major reservoirs in Canada and Japan [1, 9, 10]. Other domestic mammals, such as cats and dogs, may also play a role in the transmission to humans, for instance in Canada and Australia [11–17]. Although less frequent, birds, marine mammals, or game may be sometimes implicated in the occurrence of sporadic or clustered human cases [17–20].

This article is part of the Topical Collection on *Emerging Tropical Diseases*.

Extended author information available on the last page of the article
Considering that *C. burnetii* infection in humans is largely asymptomatic, Q fever appears to have a limited impact on public health. Nevertheless, it could cause significant disability and possibly long-term consequences due to persistent infection. Approximately 5% of those infected with *C. burnetii* develop a chronic form, now called persistent focalized *C. burnetii* infection, that can develop in patients with heart valve defects or vascular disease, immunosuppressed patients, and pregnant women. Persistent focalized Q fever can occur from a few weeks to several years after the initial infection, with patients presenting with endocarditis, pneumonia, chronic hepatitis, and other manifestations frequently less responsive to antibiotic treatment. Different frequencies of pneumonia and hepatitis have been observed depending on the regions of the world; one hypothesis is an association with local circulating strains [20–22]. Moreover, since the first descriptions in the 1990s, chronic fatigue syndrome (CFS) has been described as a sequel directly associated with *C. burnetii* infection (post-acute Q fever) [23–25]. As the clinical manifestations are nonspecific and the diagnosis complex, CFS is probably an underestimated disease that can become a major public health problem. Great difficulties are actually encountered in control, especially during and after large human outbreaks, such as in the Netherlands [26, 27], and in hyperendemic areas, such as in French Guiana [28••]. In addition, *C. burnetii* is classified as a potential inhalational bioterrorism agent [29]. These situations highlight the need for a collaborative effort from human physicians and veterinarians under the umbrella of the “one health” concept to minimize exposure among people [30–32].

In animals, infection is also usually asymptomatic [3]. The spectrum of clinical forms appears less diversified than in humans but is biased by a focus on reproductive disorders leading to substantial economic losses, known as coxiellosis. The association with late-term abortions, stillbirth, prematurity, and low birth weight is clearly demonstrated in sheep, goat, and cattle farms [33–38]. Other manifestations are suspected (infertility, metritis, mastitis) but evidence is still lacking [35]. For other animal species, knowledge on the clinical outcome of *C. burnetii* infection is very limited, but currently appears to include reproductive failure in waterbuck (*Kobus ellipsiprymnus*), roan antelope (*Hippotragus niger*), dama gazelle (*Nanger dama*), and water buffalo (*Bubalus bubalis*) and placentitis in the Pacific harbor seal (*Phoca vitulina richardsi*), steller sea lion (*Eumetopias jubatus*) and red deer (*Cervus elaphus*) [39]. Overall, knowledge is not fully understood about the pathogenesis and the sites (organs, cells) affected depending on the species. Authors agree on the circulation of *C. burnetii* strains, which appear widespread in cattle, sheep, and goats in most countries. On a technical level, the investigations were first restricted to serology.

However, the available serological tests should be optimized and validated for each species [3, 40, 41]. Since the 1990s, more and more research has included direct detection by molecular analysis (PCR). The lack of standardized methods, or at least the lack of knowledge of the performance of sensitivity and diagnostic specificity, and the limited sample size of most of the existing studies could suggest variable bias in reported prevalence rates of *C. burnetii* infection. In addition to the identification of *C. burnetii* infection in different species of animals in different regions, although some results are still questionable, it was possible to acquire descriptive knowledge such as (i) the sites and routes of excretion of ruminants such as feces, urine, vaginal secretions, semen, milk, and placenta, along with serological responses; (ii) potential environmental sources (dust, manure, etc.); and (iii) molecular epidemiological data [37, 38, 42–45]. Studies have shown a higher risk of transmission following an episode of abortion, especially in small ruminants, generating massive bacterial shedding in the environment [46]. Nevertheless, the epidemiology of *C. burnetii* infection is still complex, with particularities according to the regions of the world. A wide range of animal species can play a role in the dissemination or maintenance of the bacterium in the environment. The effective airborne propagation ranges of *C. burnetii* may vary depending on meteorological and topographical factors. It is important to better understand the diversity of epidemiological situations and evolutions around the world.

The presence of *C. burnetii* in several animal species linked to its zoonotic importance justifies the growing recent trend in scientific development related to coxiellosis [47]. The reservoirs of *C. burnetii* have been largely investigated in many countries, especially in Europe [48]. Nevertheless, the implication of wildlife in the transmission of *C. burnetii* to humans and livestock remains poorly understood [35, 48–54]. On the other hand, data on *C. burnetii* in Latin America are scarce, sparse, and extremely heterogeneous, although several studies have been published in the last decade in humans, especially in French Guiana and Brazil. On the one hand, French Guiana contains areas of hyperendemicity and holds the record for the highest annual incidence rate in the world [28••, 55]. Furthermore, there is a specific virulence associated with a pulmonary tropism, likely linked to a specific strain of the MST17 [22, 56–59]. On the other hand, countries such as Brazil have reported few cases, case series, and seroprevalence studies [60–66]. Other countries have performed only exploratory investigations, and most countries have reported none or a few human cases over the last 30 years [28••, 67]. The objective of this work is therefore to carry out a comprehensive review of the literature concerning
Considering that *C. burnetii* infection in humans is largely asymptomatic, Q fever appears to have a limited impact on public health. Nevertheless, it could cause significant disability and possibly long-term consequences due to persistent infection. Approximately 5% of those infected with *C. burnetii* develop a chronic form, now called persistent focalized *C. burnetii* infection, that can develop in patients with heart valve defects or vascular disease, immunosuppressed patients, and pregnant women. Persistent focalized Q fever can occur from a few weeks to several years after the initial infection, with patients presenting with endocarditis, pneumonia, chronic hepatitis, and other manifestations frequently less responsive to antibiotic treatment. Different frequencies of pneumonia and hepatitis have been observed depending on the regions of the world; one hypothesis is an association with local circulating strains [20–22]. Moreover, since the first descriptions in the 1990s, chronic fatigue syndrome (CFS) has been described as a sequel directly associated with *C. burnetii* infection (post-acute Q fever) [23–25]. As the clinical manifestations are nonspecific and the diagnosis complex, CFS is probably an underestimated disease that can become a major public health problem. Great difficulties are actually encountered in control, especially during and after large human outbreaks, such as in the Netherlands [26, 27], and in hyperendemic areas, such as in French Guiana [28••]. In addition, *C. burnetii* is classified as a potential inhalational bioterrorism agent [29]. These situations highlight the need for a collaborative effort from human physicians and veterinarians under the umbrella of the “one health” concept to minimize exposure among people [30–32].

In animals, infection is also usually asymptomatic [3]. The spectrum of clinical forms appears less diversified than in humans but is biased by a focus on reproductive disorders leading to substantial economic losses, known as coxiellosis. The association with late-term abortions, stillbirth, prematurity, and low birth weight is clearly demonstrated in sheep, goat, and cattle farms [33–38]. Other manifestations are suspected (infertility, metritis, mastitis) but evidence is still lacking [35]. For other animal species, knowledge on the clinical outcome of *C. burnetii* infection is very limited, but currently appears to include reproductive failure in waterbuck (*Kobus ellipsiprymnus*), roan antelope (*Hippotragus niger*), dama gazelle (*Nanger dama*), and water buffalo (*Bubalus bubalis*) and placentitis in the Pacific harbor seal (*Phoca vitulina richardsi*), steller sea lion (*Eumetopias jubatus*) and red deer (*Cervus elaphus*) [39]. Overall, knowledge is not fully understood about the pathogenesis and the sites (organs, cells) affected depending on the species. Authors agree on the circulation of *C. burnetii* strains, which appear widespread in cattle, sheep, and goats in most countries. On a technical level, the investigations were first restricted to serology.

However, the available serological tests should be optimized and validated for each species [3, 40, 41]. Since the 1990s, more and more research has included direct detection by molecular analysis (PCR). The lack of standardized methods, or at least the lack of knowledge of the performance of sensitivity and diagnostic specificity, and the limited sample size of most of the existing studies could suggest variable bias in reported prevalence rates of *C. burnetii* infection. In addition to the identification of *C. burnetii* infection in different species of animals in different regions, although some results are still questionable, it was possible to acquire descriptive knowledge such as (i) the sites and routes of excretion of ruminants such as feces, urine, vaginal secretions, semen, milk, and placenta, along with serological responses; (ii) potential environmental sources (dust, manure, etc.); and (iii) molecular epidemiological data [37, 38, 42–45]. Studies have shown a higher risk of transmission following an episode of abortion, especially in small ruminants, generating massive bacterial shedding in the environment [46]. Nevertheless, the epidemiology of *C. burnetii* infection is still complex, with particularities according to the regions of the world. A wide range of animal species can play a role in the dissemination or maintenance of the bacterium in the environment. The effective airborne propagation ranges of *C. burnetii* may vary depending on meteorological and topographical factors. It is important to better understand the diversity of epidemiological situations and evolutions around the world.

The presence of *C. burnetii* in several animal species linked to its zoonotic importance justifies the growing recent trend in scientific development related to coxiellosis [47]. The reservoirs of *C. burnetii* have been largely investigated in many countries, especially in Europe [48]. Nevertheless, the implication of wildlife in the transmission of *C. burnetii* to humans and livestock remains poorly understood [35, 48–54]. On the other hand, data on *C. burnetii* in Latin America are scarce, sparse, and extremely heterogeneous, although several studies have been published in the last decade in humans, especially in French Guiana and Brazil. On the one hand, French Guiana contains areas of hyperendemicity and holds the record for the highest annual incidence rate in the world [28••, 55]. Furthermore, there is a specific virulence associated with a pulmonary tropism, likely linked to a specific strain of the MST17 [22, 56–59]. On the other hand, countries such as Brazil have reported few cases, case series, and seroprevalence studies [60–66]. Other countries have performed only exploratory investigations, and most countries have reported none or a few human cases over the last 30 years [28••, 67]. The objective of this work is therefore to carry out a comprehensive review of the literature concerning
animal *C. burnetii* infection in Latin America and the Caribbean.

# Methodology and Objectives

## Settings

We defined Latin America as Central America, the Caribbean, and South America.

## Review of the Literature

A comprehensive literature review was carried out following the PRISMA Reporting Guidelines. The terms “Q fever” and “Coxiella” were searched together with the names of each of the 47 countries and territories of South America and the Caribbean in English, French, Spanish, and Portuguese in PubMed (National Institutes of Health’s National Library of Medicine) and ScienceDirect (Elsevier), and the terms “Coxiella” and “Q fever” in four languages in the databases SciELO (Scientific Electronic Library Online) and LILACS (Scientific Health Information from Latin America and the Caribbean countries), which are more specifically dedicated to the Latin American medical and scientific literature (Fig. 1). We excluded papers on other subjects than *C. burnetii* infection, *C. burnetii* infection in humans only, and studies on *C. burnetii* performed outside of Latin America and the Caribbean. A second time, after selecting the papers according to those inclusion criteria, the references of each

paper were analyzed independently by two authors, and the most relevant ones were also included. Lastly, unpublished data concerning *C. burnetii* according to the knowledge of the authors were also added, including, thesis, web papers, and unpublished investigations, especially in Brazil and French Guiana. The bibliography was screened for every paper, and articles without complete text but with informative abstracts or summarized in a literature review were maintained in order not to avoid some not-indexed publications, especially old ones. The research period ranged from 1 January 1950 to 31 December 2022.

The publications were then grouped and discussed into four different groups: (1) livestock (cattle, sheep, goats, equine, swine), (2) pets (mainly dogs and cats), (3) wildlife (any taxon), and (4) ticks and other ectoparasites. For each of these four parts, the results are presented in summary tables. In the second part, a special focus was made on five countries where the infection is more often reported: French Guiana, Brazil, Argentina, Chile and Colombia, and where research on animal fever has been the most abundant.

## Results/Discussion

After applying the criteria, 113 studies in French, Spanish, Portuguese or English were selected: (i) 84 published articles found in the databases; (ii) 16 articles not found in the databases but found through the bibliography review of the articles in the previous item, and (iii) 13 articles acquired from other sources, including four theses developed in Brazil and French Guiana, including four PhD theses (Fig. 1).

<figure><img src="image_3.png" /><figcaption>**Fig. 1** Flow chart of the study according to the Prisma criterion</figcaption></figure>

<figure><img src="image_4.png" /></figure>
animal *C. burnetii* infection in Latin America and the Caribbean.

# Methodology and Objectives

## Settings

We defined Latin America as Central America, the Caribbean, and South America.

## Review of the Literature

A comprehensive literature review was carried out following the PRISMA Reporting Guidelines. The terms “Q fever” and “Coxiella” were searched together with the names of each of the 47 countries and territories of South America and the Caribbean in English, French, Spanish, and Portuguese in PubMed (National Institutes of Health’s National Library of Medicine) and ScienceDirect (Elsevier), and the terms “Coxiella” and “Q fever” in four languages in the databases SciELO (Scientific Electronic Library Online) and LILACS (Scientific Health Information from Latin America and the Caribbean countries), which are more specifically dedicated to the Latin American medical and scientific literature (Fig. 1). We excluded papers on other subjects than *C. burnetii* infection, *C. burnetii* infection in humans only, and studies on *C. burnetii* performed outside of Latin America and the Caribbean. A second time, after selecting the papers according to those inclusion criteria, the references of each

paper were analyzed independently by two authors, and the most relevant ones were also included. Lastly, unpublished data concerning *C. burnetii* according to the knowledge of the authors were also added, including, thesis, web papers, and unpublished investigations, especially in Brazil and French Guiana. The bibliography was screened for every paper, and articles without complete text but with informative abstracts or summarized in a literature review were maintained in order not to avoid some not-indexed publications, especially old ones. The research period ranged from 1 January 1950 to 31 December 2022.

The publications were then grouped and discussed into four different groups: (1) livestock (cattle, sheep, goats, equine, swine), (2) pets (mainly dogs and cats), (3) wildlife (any taxon), and (4) ticks and other ectoparasites. For each of these four parts, the results are presented in summary tables. In the second part, a special focus was made on five countries where the infection is more often reported: French Guiana, Brazil, Argentina, Chile and Colombia, and where research on animal fever has been the most abundant.

## Results/Discussion

After applying the criteria, 113 studies in French, Spanish, Portuguese or English were selected: (i) 84 published articles found in the databases; (ii) 16 articles not found in the databases but found through the bibliography review of the articles in the previous item, and (iii) 13 articles acquired from other sources, including four theses developed in Brazil and French Guiana, including four PhD theses (Fig. 1).

<figure><img src="image_3.png" /><figcaption>**Fig. 1** Flow chart of the study according to the Prisma criterion</figcaption></figure>

<figure><img src="image_4.png" /></figure>
Among the 21 countries or territories in continental Latin America, 16 (76%) had at least one publication related to *C. burnetii* detection in animals, and nine (35%) among the 26 Caribbean countries and territories. The most productive country was Brazil (N=51), followed by French Guiana (N=21), Colombia (N=16), Argentina, Chile, and Uruguay (N=8), respectively. Five countries had between two and four studies, and fourteen countries or territories had only one study. About 30% of the studies have been published between 2020 and 2022. The more abundant literature was available for livestock, followed by ticks, wildlife, and at last pets.

# Animal Coxiella burnetii Infection in Latin America

## Livestock

During the study period, there were 64 publications concerning *C. burnetii* from livestock in 20 countries and territories of Latin America and the Caribbean, mainly from Brazil (N=18), Colombia (N=10), and French Guiana (N=6) (Fig. 2). Most of them concerned cattle, sheep, and goats, but also included zebu cattle, buffalo cattle, alpaca, horses, swine, and chicken (Table 1). Most of the studies were seroprevalence surveys in livestock, especially cattle. In cattle, seroprevalence rates varied a lot according to regions and studies, from 0% in the

Brazilian state of Mato Grosso do Sul and in several Islands from the lesser Caribbean to 31.8% in Duque de Caxias, state of Rio de Janeiro [72, 83, 101]. Some authors also calculated the herd prevalence in addition to the apparent individual prevalence, with a variation from 46.9 to 82% [102, 105, 122]. Seroprevalence in goats varied a lot, from 0%, in the Caribbean Islands, in French Guiana or Piauí state, Brazil, to 35% in Nuevo Leon, Mexico, 50% in Itaboraí, Brazilian state of Rio de Janeiro, and 60.6% in Venezuela [76, 78, 101, 107, 109, 111, 121]. In most of the studies, the seroprevalence was relatively low, between 0 and 3% [80, 98, 100, 101]. At last, seroprevalence in sheep was highly variable also, from 0% in Argentina and the Caribbean Islands Trinidad and French Guiana, to 17.3% in Uruguay, 26.3% in St. Kitts, and 66% in Itaboraí, Brazilian state of Rio de Janeiro [76, 113–116].

Some had investigated the presence of the *C. burnetii* genome on vaginal swabs [86, 89, 98, 109], fetuses or placenta [86, 88••, 89, 117], serum or blood samples [66, 76, 99], and finally, milk and dairy products [71, 76, 81, 84, 85, 96, 98, 123]. Vaginal carriage of *C. burnetii* has been found in a few studies and implicated as a cause of abortion for different species [63, 88••, 118–120].

*C. burnetii* infection has been reported elsewhere in other domestic mammals, though less frequently than in livestock [124]. In Latin America, studies among domestic animals, including cattle (including zebu cattle and buffalo cattle),

<figure><img src="image_5.png" /><figcaption>**Fig. 2** Map of the publications on *Coxiella burnetii* infection in livestock in Latin America and the Caribbean</figcaption></figure>
Among the 21 countries or territories in continental Latin America, 16 (76%) had at least one publication related to *C. burnetii* detection in animals, and nine (35%) among the 26 Caribbean countries and territories. The most productive country was Brazil (N=51), followed by French Guiana (N=21), Colombia (N=16), Argentina, Chile, and Uruguay (N=8), respectively. Five countries had between two and four studies, and fourteen countries or territories had only one study. About 30% of the studies have been published between 2020 and 2022. The more abundant literature was available for livestock, followed by ticks, wildlife, and at last pets.

# Animal Coxiella burnetii Infection in Latin America

## Livestock

During the study period, there were 64 publications concerning *C. burnetii* from livestock in 20 countries and territories of Latin America and the Caribbean, mainly from Brazil (N=18), Colombia (N=10), and French Guiana (N=6) (Fig. 2). Most of them concerned cattle, sheep, and goats, but also included zebu cattle, buffalo cattle, alpaca, horses, swine, and chicken (Table 1). Most of the studies were seroprevalence surveys in livestock, especially cattle. In cattle, seroprevalence rates varied a lot according to regions and studies, from 0% in the

Brazilian state of Mato Grosso do Sul and in several Islands from the lesser Caribbean to 31.8% in Duque de Caxias, state of Rio de Janeiro [72, 83, 101]. Some authors also calculated the herd prevalence in addition to the apparent individual prevalence, with a variation from 46.9 to 82% [102, 105, 122]. Seroprevalence in goats varied a lot, from 0%, in the Caribbean Islands, in French Guiana or Piauí state, Brazil, to 35% in Nuevo Leon, Mexico, 50% in Itaboraí, Brazilian state of Rio de Janeiro, and 60.6% in Venezuela [76, 78, 101, 107, 109, 111, 121]. In most of the studies, the seroprevalence was relatively low, between 0 and 3% [80, 98, 100, 101]. At last, seroprevalence in sheep was highly variable also, from 0% in Argentina and the Caribbean Islands Trinidad and French Guiana, to 17.3% in Uruguay, 26.3% in St. Kitts, and 66% in Itaboraí, Brazilian state of Rio de Janeiro [76, 113–116].

Some had investigated the presence of the *C. burnetii* genome on vaginal swabs [86, 89, 98, 109], fetuses or placenta [86, 88••, 89, 117], serum or blood samples [66, 76, 99], and finally, milk and dairy products [71, 76, 81, 84, 85, 96, 98, 123]. Vaginal carriage of *C. burnetii* has been found in a few studies and implicated as a cause of abortion for different species [63, 88••, 118–120].

*C. burnetii* infection has been reported elsewhere in other domestic mammals, though less frequently than in livestock [124]. In Latin America, studies among domestic animals, including cattle (including zebu cattle and buffalo cattle),

<figure><img src="image_5.png" /><figcaption>**Fig. 2** Map of the publications on *Coxiella burnetii* infection in livestock in Latin America and the Caribbean</figcaption></figure>
<table><caption>**Table 1** Data on *Coxiella burnetii* detection in livestock, swine and equine in Latin America</caption><thead><tr><th>Author</th><th>Year of the study</th><th>Year of publication</th><th>Language</th><th>Country</th><th>State</th><th>Methodology</th><th>Sample type</th><th>Species</th><th>Number of samples</th><th>Result</th><th>Comments</th><th>Publication in a journal</th></tr></thead><tbody><tr><td>Babudieri and Kaplan [68, 69]</td><td>NA</td><td>1952</td><td>Spanish</td><td>Argentina</td><td>–</td><td>CFT</td><td>Serum</td><td>Sheep</td><td>0/22</td><td>0%</td><td></td><td>Yes</td></tr><tr><td>Trezeguet [70]</td><td>2007</td><td>2010</td><td>Spanish</td><td>Argentina</td><td>Various states</td><td>ELISA and CFT</td><td>Serum</td><td>Goats</td><td>33/15,272</td><td>0.22%</td><td></td><td>Yes</td></tr><tr><td>Rojas [71]</td><td>NA</td><td>2013</td><td>Spanish</td><td>Argentina</td><td>NA</td><td>RT-PCR (IS1111)</td><td>Milk</td><td>Cattle</td><td>0/1</td><td>0%</td><td></td><td>Yes</td></tr><tr><td>Travassos and Kaplan [72, 68]</td><td>NA</td><td>1954</td><td>Portuguese</td><td>Brazil</td><td>Caixas (vicinity of Rio de Janeiro)</td><td>CFT</td><td>Serum</td><td>Cattle</td><td>114/358</td><td>31.8%</td><td></td><td>Yes</td></tr><tr><td>Valle [73]</td><td>NA</td><td>1955</td><td>Portuguese</td><td>Brazil</td><td>São Paulo</td><td>CFT</td><td>Serum</td><td>Cattle</td><td>24/171</td><td>14%</td><td></td><td>Yes</td></tr><tr><td>Riemann [74]</td><td>1972</td><td>1975</td><td>English</td><td>Brazil</td><td>7 towns of Minas Gerais</td><td>IHA</td><td>Serum</td><td>Zebu cattle</td><td>45/156</td><td>29%</td><td></td><td>Yes</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td>State and 12 towns of Goiás</td><td></td><td></td><td></td><td></td><td></td><td>Study in slaughter-house</td><td></td></tr><tr><td>Brown [75]</td><td>NA</td><td>1989</td><td>English</td><td>Brazil</td><td>5 among 9 States in North-east regions</td><td>ELISA</td><td>Serum</td><td>Goats</td><td>0/76</td><td>0%</td><td></td><td>Yes</td></tr><tr><td>Mares-Guia [76]</td><td>2008–2009</td><td>2014</td><td>English</td><td>Brazil</td><td>Itaboraí municipality, State of Rio de Janeiro</td><td>IFA</td><td>Serum</td><td>Sheep</td><td>2/3</td><td>66.6%</td><td></td><td>Yes</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td>PCR (httpAB gene)</td><td>Serum</td><td>Sheep</td><td>0/2</td><td>0%</td><td></td><td>Area of the first human case of Q fever in Brazil</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Milk</td><td>Goats</td><td>5/10</td><td>50%</td><td></td><td></td></tr><tr><td>Mares-Guia [77]</td><td>2011–2012</td><td>2015</td><td>Portuguese</td><td>Brazil</td><td>Itaboraí municipality, State of Rio de Janeiro</td><td>IFA</td><td>Serum</td><td>Sheep</td><td>2/15</td><td>17.1%</td><td></td><td>Several human cases in the area</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td>PCR (httpAB gene)</td><td>Serum</td><td>Sheep</td><td>0/15</td><td>0%</td><td></td><td>Unpublished thesis</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Udder</td><td>Sheep</td><td>1/6</td><td>16.7%</td><td></td><td></td></tr></tbody></table>

<figure><img src="image_6.png" /></figure>
<table><caption>**Table 1** Data on *Coxiella burnetii* detection in livestock, swine and equine in Latin America</caption><thead><tr><th>Author</th><th>Year of the study</th><th>Year of publication</th><th>Language</th><th>Country</th><th>State</th><th>Methodology</th><th>Sample type</th><th>Species</th><th>Number of samples</th><th>Result</th><th>Comments</th><th>Publication in a journal</th></tr></thead><tbody><tr><td>Babudieri and Kaplan [68, 69]</td><td>NA</td><td>1952</td><td>Spanish</td><td>Argentina</td><td>–</td><td>CFT</td><td>Serum</td><td>Sheep</td><td>0/22</td><td>0%</td><td></td><td>Yes</td></tr><tr><td>Trezeguet [70]</td><td>2007</td><td>2010</td><td>Spanish</td><td>Argentina</td><td>Various states</td><td>ELISA and CFT</td><td>Serum</td><td>Goats</td><td>33/15,272</td><td>0.22%</td><td></td><td>Yes</td></tr><tr><td>Rojas [71]</td><td>NA</td><td>2013</td><td>Spanish</td><td>Argentina</td><td>NA</td><td>RT-PCR (IS1111)</td><td>Milk</td><td>Cattle</td><td>0/1</td><td>0%</td><td></td><td>Yes</td></tr><tr><td>Travassos and Kaplan [72, 68]</td><td>NA</td><td>1954</td><td>Portuguese</td><td>Brazil</td><td>Caixas (vicinity of Rio de Janeiro)</td><td>CFT</td><td>Serum</td><td>Cattle</td><td>114/358</td><td>31.8%</td><td></td><td>Yes</td></tr><tr><td>Valle [73]</td><td>NA</td><td>1955</td><td>Portuguese</td><td>Brazil</td><td>São Paulo</td><td>CFT</td><td>Serum</td><td>Cattle</td><td>24/171</td><td>14%</td><td></td><td>Yes</td></tr><tr><td>Riemann [74]</td><td>1972</td><td>1975</td><td>English</td><td>Brazil</td><td>7 towns of Minas Gerais State and 12 towns of Goiás State</td><td>IHA</td><td>Serum</td><td>Zebu cattle</td><td>45/156</td><td>29%</td><td>Study in slaughter-house</td><td>Yes</td></tr><tr><td>Brown [75]</td><td>NA</td><td>1989</td><td>English</td><td>Brazil</td><td>5 among 9 States in North-east regions</td><td>ELISA</td><td>Serum</td><td>Goats</td><td>0/76</td><td>0%</td><td></td><td>Yes</td></tr><tr><td>Mares-Guia [76]</td><td>2008–2009</td><td>2014</td><td>English</td><td>Brazil</td><td>Itaboraí municipality, State of Rio de Janeiro</td><td>IFA</td><td>Serum</td><td>Sheep</td><td>2/3</td><td>66.6%</td><td>Area of the first human case of Q fever in Brazil</td><td>Yes</td></tr><tr><td>Mares-Guia [77]</td><td>2011–2012</td><td>2015</td><td>Portuguese</td><td>Brazil</td><td>Itaboraí municipality, State of Rio de Janeiro</td><td>PCR (httpAB gene)</td><td>Serum</td><td>Sheep</td><td>0/2</td><td>0%</td><td></td><td>Yes</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td>IFA</td><td>Serum</td><td>Sheep</td><td>2/15</td><td>17.1%</td><td>Several human cases in the area</td><td>Yes</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td>PCR (httpAB gene)</td><td>Serum</td><td>Sheep</td><td>0/15</td><td>0%</td><td></td><td>Unpublished thesis</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Udder</td><td>Sheep</td><td>1/6</td><td>16.7%</td><td></td><td></td></tr></tbody></table>

<figure><img src="image_6.png" /></figure>
<table><caption>Table 1 (continued)</caption><thead><tr><th>Author</th><th>Year of the study</th><th>Year of publication</th><th>Language</th><th>Country</th><th>State</th><th>Methodology</th><th>Sample type</th><th>Species</th><th>Number of samples</th><th>Result</th><th>Comments</th><th>Publication in a journal</th></tr></thead><tbody><tr><td>Guimarães [78]</td><td>2013</td><td>2017</td><td>Portuguese</td><td>Brazil</td><td>Parque Nacional da Serra das Confluentes, Caatinga biome, Piauí</td><td>IFA</td><td>Serum</td><td>Sheep</td><td>3/153</td><td>2.0%</td><td>Not on Pub-Med</td><td>Yes</td></tr><tr><td>De Oliveira [79]</td><td>2014–2015</td><td>2018</td><td>English</td><td>Brazil</td><td>Agreste mesoregion, Alagoas</td><td>ELISA PCR (IS1111)</td><td>Serum Placenta</td><td>Goats</td><td>0/202</td><td>0%</td><td></td><td></td></tr><tr><td>Souza [80]</td><td>2016</td><td>2018</td><td>English</td><td>Brazil</td><td>Petrolina, Pernambuco State</td><td>IFA</td><td>Serum</td><td>Goats</td><td>9/412</td><td>2.2%</td><td>Dairy goats with reproductive disorders</td><td>Yes</td></tr><tr><td>Mioni [81]</td><td>2017</td><td>2019</td><td>English</td><td>Brazil</td><td>Goiás State</td><td>qPCR (IS1111)</td><td>Raw milk</td><td>Cattle</td><td>4/112</td><td>3.6%</td><td></td><td>Yes</td></tr><tr><td>Zanatto [82]</td><td>2014–2015</td><td>2019</td><td>English</td><td>Brazil</td><td>States São Paulo, Minas Gerais Minas, Mato Grosso do Sul, Goiás</td><td>IFA</td><td>Serum</td><td>Cattle</td><td>14/102</td><td>13.7%</td><td>Cows showing a history of reproductive disorders</td><td>Yes</td></tr><tr><td>Ramos [83]</td><td>NA</td><td>2020</td><td>English</td><td>Brazil</td><td>Mato Grosso do Sul</td><td>IFA</td><td>Serum</td><td>Zebu cattle</td><td></td><td></td><td></td><td>Yes</td></tr><tr><td>Rozental [84, 85]</td><td>NA</td><td>2020</td><td>English</td><td>Brazil</td><td>Serro microregion, Minas Gerais State</td><td>PCR (IS1111)</td><td>Artisanal cheese</td><td>Cow Calves</td><td>2/200</td><td>1.0%</td><td></td><td>Yes</td></tr><tr><td>Mioni [86]</td><td>NA</td><td>2020</td><td>English</td><td>Brazil</td><td>São Paulo</td><td>Genotyping (MLVA and MST) #</td><td>Vaginal swabs</td><td>Cattle</td><td>5/53</td><td>9.4%</td><td></td><td>Yes</td></tr><tr><td>Mioni [64]</td><td>2014–2015</td><td>2020</td><td>English</td><td>Brazil</td><td>São Paulo State</td><td>IFA qPCR (IS1111)</td><td>Fetus Serum</td><td>Cattle and Zebu cattle</td><td>3/3</td><td>100%</td><td>MST74</td><td>Yes</td></tr></tbody></table>

<figure><img src="image_7.png" /></figure>
<table><caption>Table 1 (continued)</caption><thead><tr><th>Author</th><th>Year of the study</th><th>Year of publication</th><th>Language</th><th>Country</th><th>State</th><th>Methodology</th><th>Sample type</th><th>Species</th><th>Number of samples</th><th>Result</th><th>Comments</th><th>Publication in a journal</th></tr></thead><tbody><tr><td>Guimarães [78]</td><td>2013</td><td>2017</td><td>Portuguese</td><td>Brazil</td><td>Parque Nacional da Serra das Confluentes, Caatinga biome, Piauí</td><td>IFA</td><td>Serum</td><td>Sheep</td><td>3/153</td><td>2.0%</td><td>Not on Pub-Med</td><td>Yes</td></tr><tr><td>De Oliveira [79]</td><td>2014–2015</td><td>2018</td><td>English</td><td>Brazil</td><td>Agreste mesoregion, Alagoas</td><td>ELISA PCR (IS1111)</td><td>Serum Placenta</td><td>Goats</td><td>0/202</td><td>0%</td><td></td><td></td></tr><tr><td>Souza [80]</td><td>2016</td><td>2018</td><td>English</td><td>Brazil</td><td>Petrolina, Pernambuco State</td><td>IFA</td><td>Serum</td><td>Goats</td><td>172/312</td><td>55.1%</td><td>Dairy goats with reproductive disorders</td><td>Yes</td></tr><tr><td>Mioni [81]</td><td>2017</td><td>2019</td><td>English</td><td>Brazil</td><td>Goiás State</td><td>qPCR (IS1111)</td><td>Raw milk</td><td>Cattle</td><td>9/412</td><td>2.2%</td><td></td><td></td></tr><tr><td>Zanatto [82]</td><td>2014–2015</td><td>2019</td><td>English</td><td>Brazil</td><td>States São Paulo, Minas Gerais Minas, Mato Grosso do Sul, Goiás</td><td>IFA</td><td>Serum</td><td>Cattle</td><td>9/403</td><td>2.1%</td><td></td><td></td></tr><tr><td>Ramos [83]</td><td>NA</td><td>2020</td><td>English</td><td>Brazil</td><td>Mato Grosso do Sul</td><td>IFA</td><td>Serum</td><td>Zebu cattle</td><td>2/200</td><td>1.0%</td><td></td><td></td></tr><tr><td>Rozental [84, 85]</td><td>NA</td><td>2020</td><td>English</td><td>Brazil</td><td>Serro microregion, Minas Gerais State</td><td>PCR (IS1111)</td><td>Artisanal cheese</td><td>Cow Calves</td><td>0/200</td><td>0%</td><td></td><td></td></tr><tr><td>Mioni [86]</td><td>NA</td><td>2020</td><td>English</td><td>Brazil</td><td>São Paulo</td><td>Genotyping (MLVA and MST) #</td><td>Vaginal swabs</td><td>Sheep</td><td>3/3</td><td>100%</td><td>MST74</td><td>Yes</td></tr><tr><td>Mioni [64]</td><td>2014–2015</td><td>2020</td><td>English</td><td>Brazil</td><td>São Paulo State</td><td>IFA qPCR (IS1111)</td><td>Fetus Serum</td><td>Goat</td><td>1/1</td><td>100%</td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Cattle</td><td>3/3</td><td>100%</td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Cattle and Zebu cattle</td><td>360/1515</td><td>23.8%</td><td>45/54; 83.3%</td><td>Yes of the cities had at least 1 seropositive animal</td></tr></tbody></table>

<figure><img src="image_7.png" /></figure>
<table><caption>Table 1 (continued)</caption><thead><tr><th>Author</th><th>Year of the study</th><th>Year of publication</th><th>Language</th><th>Country</th><th>State</th><th>Methodology</th><th>Sample type</th><th>Species</th><th>Number of samples</th><th>Result</th><th>Comments</th><th>Publication in a journal</th></tr></thead><tbody><tr><td>Nascimento [87]</td><td>2020</td><td>2021</td><td>English</td><td>Brazil</td><td>Municipality of Patos de Minas, Cerrado, Mineiro, Minas Gerais State</td><td>qPCR (IS1111)</td><td>Artisanal cheese</td><td>NA (likely cattle)</td><td>4/87</td><td>4.6%</td><td>Yes</td><td>Yes</td></tr><tr><td>Mioni [88••]</td><td>2013–2019</td><td>2022</td><td>English</td><td>Brazil</td><td>São Paulo State University and Universidade Federal do Rio Grande do Sul</td><td></td><td>Fetuses</td><td>Cattle</td><td>7/76</td><td>9.2%</td><td>3 co-infec-tions with Neospora spp.</td><td></td></tr><tr><td>Mioni [89]</td><td>NA</td><td>Unpublished</td><td>Portuguese</td><td>Brazil</td><td>São Paulo</td><td>qPCR IS1111</td><td>Vaginal swabs</td><td>Sheep and goats</td><td>27/133</td><td>20.3%</td><td></td><td></td></tr><tr><td>International Society for Infectious Diseases [90]</td><td>2014</td><td>2014</td><td>English</td><td>Chile</td><td>NA</td><td>ELISA</td><td>Serum</td><td>Alpaca (Vicugna pacos)</td><td>13/946</td><td>1.4%</td><td></td><td></td></tr><tr><td>Cornejo [91]</td><td>2017</td><td>2019</td><td>English</td><td>Chile</td><td>Metropolitan, Nuble, Bío-Bío, Araucania, Los Ríos, and Los Lagos Districts</td><td>qPCR IS1111</td><td>Raw tank milk from 105 different farms</td><td>Cattle</td><td>2/105</td><td>2.1%</td><td></td><td></td></tr></tbody></table>

<figure><img src="image_8.png" /></figure>
<table><caption>Table 1 (continued)</caption><thead><tr><th>Author</th><th>Year of the study</th><th>Year of publication</th><th>Language</th><th>Country</th><th>State</th><th>Methodology</th><th>Sample type</th><th>Species</th><th>Number of samples</th><th>Result</th><th>Comments</th><th>Publication in a journal</th></tr></thead><tbody><tr><td>Nascimento [87]</td><td>2020</td><td>2021</td><td>English</td><td>Brazil</td><td>Municipality of Patos de Minas, Cerrado, Mineiro, Minas Gerais State</td><td>qPCR (IS1111)</td><td>Artisanal cheese</td><td>NA (likely cattle)</td><td>4/87</td><td>4.6%</td><td>Yes</td><td>Yes</td></tr><tr><td>Mioni [88••]</td><td>2013–2019</td><td>2022</td><td>English</td><td>Brazil</td><td>São Paulo State University and Universidade Federal do Rio Grande do Sul</td><td></td><td>Fetuses</td><td>Cattle</td><td>7/76</td><td>9.2%</td><td>3 co-infec- tions with Neospora spp.</td><td></td></tr><tr><td>Mioni [89]</td><td>NA</td><td>Unpublished</td><td>Portuguese</td><td>Brazil</td><td>São Paulo State University and Universidade Federal do Rio Grande do Sul</td><td>qPCR IS1111</td><td>Vaginal swabs</td><td>Sheep and goats</td><td>27/133</td><td>20.3%</td><td>Sheep: flock with the reproductive disorder (abortions)</td><td>No</td></tr><tr><td>International Society for Infectious Diseases [90]</td><td>2014</td><td>2014</td><td>English</td><td>Chile</td><td></td><td>ELISA</td><td>Serum</td><td>Alpaca (Vicugna pacos)</td><td>13/946</td><td>1.4%</td><td>Goats: asymptomatic flock</td><td>No</td></tr><tr><td>Cornejo [91]</td><td>2017</td><td>2019</td><td>English</td><td>Chile</td><td>Metropolitan, Nuble, Bío-Bío, Araucania, Los Ríos, and Los Lagos Districts</td><td>qPCR IS1111</td><td>Raw tank milk from 105 different farms</td><td>Cattle</td><td>2/105</td><td>2.1%</td><td>Culled after importation in China after being diagnosed with Q fever</td><td>Yes</td></tr></tbody></table>

<figure><img src="image_8.png" /></figure>
<table><caption>**Table 1 (continued)**</caption><thead><tr><th>Author</th><th>Year of the study</th><th>Year of publication</th><th>Language</th><th>Country</th><th>State</th><th>Methodology</th><th>Sample type</th><th>Species</th><th>Number of samples</th><th>Result</th><th>Comments</th><th>Publication in a journal</th></tr></thead><tbody><tr><td>Covelli [92]</td><td>NA</td><td>1961</td><td>Spanish</td><td>Colombia</td><td>Departamentos de Antioquia, Cundinamarca and Meta</td><td>CTF</td><td>Serum</td><td>Cattle</td><td>8/1715</td><td>0.46%</td><td></td><td>Yes</td></tr><tr><td>San Martin-Berberi [93]</td><td>1959</td><td>1963</td><td>Spanish</td><td>Colombia</td><td>Valle, Cali</td><td>MAT</td><td>Milk</td><td></td><td></td><td></td><td>"Proof of infection"</td><td>No personal data</td></tr><tr><td>Vaughn [93]</td><td>1963</td><td>1967</td><td>Spanish</td><td>Colombia</td><td>Valle, Caldas, Cauca, Tolima, Nariño, Santander, Caquetá, Cundinamarca, and Huila</td><td>MAT</td><td>Serum</td><td>Cattle</td><td>110/454</td><td>24.2%</td><td>Animals from slaughterhouse</td><td>Yes</td></tr><tr><td>Lorbacher de Ruiz [94]</td><td>NA</td><td>1977</td><td>German</td><td>Colombia</td><td>Various</td><td>CFT</td><td>Serum</td><td>Horses Cattle</td><td>89/357</td><td>25.0%</td><td>Manuscript not available. Data in a review of the literature</td><td>Yes</td></tr><tr><td>Perry [95]</td><td>NA</td><td>1981</td><td>Spanish</td><td>Colombia</td><td>Boyacá Department</td><td>CFT</td><td>Serum</td><td>Sheep Imported Local</td><td>197/2501 91/402 106/2099</td><td>7.9% 23.0% 5.0%</td><td>Blackface sheep recently imported from Great Britain</td><td>Yes</td></tr><tr><td>Rojas [71]</td><td>NA</td><td>2013</td><td>Spanish</td><td>Colombia</td><td>NA</td><td>RT-PCR (IS1111)</td><td>Milk</td><td>Cattle</td><td>1/4</td><td>25%</td><td>Ct 37.32</td><td>Yes</td></tr><tr><td>Contreras [96]</td><td>2012</td><td>2015</td><td>English</td><td>Colombia</td><td>Montería</td><td>IFA qPCR (IS1111)</td><td>Milk</td><td>Cattle</td><td>5/11</td><td>45%</td><td>Strain CbuK 154Q</td><td>Yes</td></tr><tr><td>Eraso-Cadena [97]</td><td>2014</td><td>2017</td><td>English</td><td>Colombia</td><td>North and Magdalena Medio subregions of Antioquia</td><td>IFA</td><td>Serum</td><td>Cattle</td><td>104/384</td><td>27.1%</td><td>48 livestock farms</td><td>Yes</td></tr></tbody></table>

<figure><img src="image_9.png" /></figure>
<table><caption>**Table 1 (continued)**</caption><thead><tr><th>Author</th><th>Year of the study</th><th>Year of publication</th><th>Language</th><th>Country</th><th>State</th><th>Methodology</th><th>Sample type</th><th>Species</th><th>Number of samples</th><th>Result</th><th>Comments</th><th>Publication in a journal</th></tr></thead><tbody><tr><td>Covelli [92]</td><td>NA</td><td>1961</td><td>Spanish</td><td>Colombia</td><td>Departamentos de Antioquia, Cundinamarca and Meta</td><td>CTF</td><td>Serum</td><td>Cattle</td><td>8/1715</td><td>0.46%</td><td></td><td>Yes</td></tr><tr><td>San Martin-Berberi [93]</td><td>1959</td><td>1963</td><td>Spanish</td><td>Colombia</td><td>Valle, Cali</td><td>MAT</td><td>Milk</td><td></td><td></td><td></td><td>"Proof of infection"</td><td>No personal data</td></tr><tr><td>Vaughn [93]</td><td>1963</td><td>1967</td><td>Spanish</td><td>Colombia</td><td>Valle, Caldas, Cauca, Tolima, Nariño, Santander, Caquetá, Cundinamarca, and Huila</td><td>MAT</td><td>Serum</td><td>Cattle</td><td>110/454</td><td>24.2%</td><td>Animals from slaughterhouse</td><td>Yes</td></tr><tr><td>Lorbacher de Ruiz [94]</td><td>NA</td><td>1977</td><td>German</td><td>Colombia</td><td>Various</td><td>CFT</td><td>Serum</td><td>Horses Cattle</td><td>89/357</td><td>25.0%</td><td>Manuscript not available. Data in a review of the literature</td><td>Yes</td></tr><tr><td>Perry [95]</td><td>NA</td><td>1981</td><td>Spanish</td><td>Colombia</td><td>Boyacá Department</td><td>CFT</td><td>Serum</td><td>Sheep Imported Local</td><td>197/2501 91/402 106/2099</td><td>7.9% 23.0% 5.0%</td><td>Blackface sheep recently imported from Great Britain</td><td>Yes</td></tr><tr><td>Rojas [71]</td><td>NA</td><td>2013</td><td>Spanish</td><td>Colombia</td><td>NA</td><td>RT-PCR (IS1111)</td><td>Milk</td><td>Cattle</td><td>1/4</td><td>25%</td><td>Ct 37.32</td><td>Yes</td></tr><tr><td>Contreras [96]</td><td>2012</td><td>2015</td><td>English</td><td>Colombia</td><td>Montería</td><td>IFA qPCR (IS1111)</td><td>Milk</td><td>Cattle</td><td>5/11</td><td>45%</td><td>Strain CbuK 154Q</td><td>Yes</td></tr><tr><td>Eraso-Cadena [97]</td><td>2014</td><td>2017</td><td>English</td><td>Colombia</td><td>North and Magdalena Medio subregions of Antioquia</td><td>IFA</td><td>Serum</td><td>Cattle</td><td>104/384</td><td>27.1%</td><td>48 livestock farms</td><td>Yes</td></tr></tbody></table>

<figure><img src="image_9.png" /></figure>
<table><caption>**Table 1 (continued)**</caption><thead><tr><th>Author</th><th>Year of the study</th><th>Year of publication</th><th>Language</th><th>Country</th><th>State</th><th>Methodology</th><th>Sample type</th><th>Species</th><th>Number of samples</th><th>Result</th><th>Comments</th><th>Publication in a journal</th></tr></thead><tbody><tr><td>Contreras [98]</td><td>2013</td><td>2018</td><td>English</td><td>Colombia</td><td>Municipality of Valledupar, department of Cesar</td><td>Conventional PCR (IS1111)</td><td>Milk</td><td>Sheep</td><td>4/66</td><td>6%</td><td>Percentage of identity 100% strain CbuK Q_154</td><td></td></tr><tr><td>Cabrera Orrego [99]</td><td>2015</td><td>2020</td><td>English</td><td>Colombia</td><td>Magdalena Medio region, Antioquia</td><td>qPCR (IS 1111) 16SRNA</td><td>Vaginal swabs</td><td>Goats</td><td>2/328</td><td>0.6%</td><td>99% strain Guiana Cb175</td><td></td></tr><tr><td>Villagra-Blanc [100]</td><td>2013–2017</td><td>2018</td><td>English</td><td>Costa Rica</td><td>Central, North Huetar, Atlantic Huetar, Central Pacific, Chorotega, Brunca</td><td>ELISA</td><td>Serum</td><td>Goats Flock</td><td>7/391 1/13</td><td>1.8% 7.7%</td><td>Risk factors: Location: Municipality (Puerto Nare), female, born on the farm, intended for meat production, spent over 49 months of residence on the farm</td><td>Yes</td></tr><tr><td>Johnson [101]</td><td>NA</td><td>2020</td><td>English</td><td>Dominica</td><td>NA</td><td>IFA</td><td>Serum</td><td>Cattle Sheep Goats</td><td>0/83 1/54 0/65</td><td>0% 2% 0%</td><td>Detected only in 1 flock in the North Huetar Region. 15.2% of the goat positive in this flock</td><td>Yes</td></tr></tbody></table>

<figure><img src="image_10.png" /></figure>
<table><caption>**Table 1 (continued)**</caption><thead><tr><th>Author</th><th>Year of the study</th><th>Year of publication</th><th>Language</th><th>Country</th><th>State</th><th>Methodology</th><th>Sample type</th><th>Species</th><th>Number of samples</th><th>Result</th><th>Comments</th><th>Publication in a journal</th></tr></thead><tbody><tr><td>Contreras [98]</td><td>2013</td><td>2018</td><td>English</td><td>Colombia</td><td>Municipality of Valledupar, department of Cesar</td><td>Conventional PCR (IS1111)</td><td>Milk</td><td>Sheep</td><td>4/66</td><td>6%</td><td>Percentage of identity 100% strain CbuK Q_154</td><td></td></tr><tr><td>Cabrera Orrego [99]</td><td>2015</td><td>2020</td><td>English</td><td>Colombia</td><td>Magdalena Medio region, Antioquia</td><td>qPCR (IS 1111) 16SRNA</td><td>Vaginal swabs</td><td>Goats</td><td>2/328</td><td>0.6%</td><td>99% strain Guiana Cb175</td><td></td></tr><tr><td>Villagra-Blanc [100]</td><td>2013–2017</td><td>2018</td><td>English</td><td>Costa Rica</td><td>Central, North Huetar, Atlantic Huetar, Central Pacific, Chorotega, Brunca</td><td>ELISA</td><td>Serum</td><td>Goats Flock</td><td>7/391 1/13</td><td>1.8% 7.7%</td><td>Risk factors: Location: Municipality (Puerto Nare), female, born on the farm, intended for meat production, spent over 49 months of residence on the farm</td><td>Yes</td></tr><tr><td>Johnson [101]</td><td>NA</td><td>2020</td><td>English</td><td>Dominica</td><td>NA</td><td>IFA</td><td>Serum</td><td>Cattle Sheep Goats</td><td>0/83 1/54 0/65</td><td>0% 2% 0%</td><td>Detected only in 1 flock in the North Huetar Region. 15.2% of the goat positive in this flock</td><td>Yes</td></tr></tbody></table>

<figure><img src="image_10.png" /></figure>
<table><caption>**Table 1 (continued)**</caption><thead><tr><th>Author</th><th>Year of the study</th><th>Year of publication</th><th>Language</th><th>Country</th><th>State</th><th>Methodology</th><th>Sample type</th><th>Species</th><th>Number of samples</th><th>Result</th><th>Comments</th><th>Publication in a journal</th></tr></thead><tbody><tr><td>Rojas [71]</td><td>NA</td><td>2013</td><td>Spanish</td><td>Ecuador</td><td>Cruda, Cotopaxi, Carchi, Loja, Chimborazo, Pastaza, Guayas, Galápagos, Imabura</td><td>RT-PCR (IS111)</td><td>Milk</td><td>Cattle</td><td>3/102</td><td>2.9%</td><td>All in the Chimborazo province<br>Ct: 36.62, 32.73, and 37.10</td><td>Yes</td></tr><tr><td>Carbonero [102]</td><td>2008–2010</td><td>2015</td><td>English</td><td>Ecuador</td><td>Azuay, Chimborazo, Cotopaxi, Manabí, Pichincha, Santo Domingo, Tungurahua and Zamora-Chinchipe</td><td>ELISA</td><td>Serum</td><td>Cattle<br>Apparent individual prevalence</td><td>386/2668</td><td>14.5%</td><td>2668 serum samples from 386 herds</td><td>Risk factors: females (15.5 vs. 8.5%), cattle > 4 years old (20.2 vs. 10.6%)</td></tr><tr><td>Echeverria [103]</td><td>NA</td><td>2019</td><td>English</td><td>Ecuador</td><td>Quito area (Andes region) and Santo Domingo de los Tsachilas (tropical region)</td><td>ELISA</td><td>Serum</td><td>Cattle</td><td>151/352</td><td>42.9%</td><td>Dairy cattle</td><td></td></tr><tr><td>Changoluisa [104]</td><td>NA</td><td>2019</td><td>English</td><td>Ecuador</td><td>Santo Domingo Province</td><td>ELISA</td><td>Serum</td><td>Cattle<br>Cows with a first-calf abortion</td><td>91/172<br>47/95</td><td>52.9%<br>49.4%</td><td></td><td>Yes</td></tr><tr><td>Rice [105]</td><td>NA</td><td>1979</td><td>English</td><td>El Salvador</td><td>NA</td><td>CFT</td><td>Serum</td><td>Control cows<br>Cattle<br>Positive farms</td><td>44/77<br>77/297<br>27/33</td><td>57%<br>26.0%<br>82.0%</td><td></td><td>Yes</td></tr></tbody></table>
<table><caption>**Table 1 (continued)**</caption><thead><tr><th>Author</th><th>Year of the study</th><th>Year of publication</th><th>Language</th><th>Country</th><th>State</th><th>Methodology</th><th>Sample type</th><th>Species</th><th>Number of samples</th><th>Result</th><th>Comments</th><th>Publication in a journal</th></tr></thead><tbody><tr><td>Rojas [71]</td><td>NA</td><td>2013</td><td>Spanish</td><td>Ecuador</td><td>Cruda, Cotopaxi, Carchi, Loja, Chimborazo, Pastaza, Guayas, Galápagos, Imabura</td><td>RT-PCR (IS111)</td><td>Milk</td><td>Cattle</td><td>3/102</td><td>2.9%</td><td>All in the Chimborazo province<br>Ct: 36.62, 32.73, and 37.10</td><td>Yes</td></tr><tr><td>Carbonero [102]</td><td>2008–2010</td><td>2015</td><td>English</td><td>Ecuador</td><td>Azuay, Chimborazo, Cotopaxi, Manabí, Pichincha, Santo Domingo, Tungurahua and Zamora-Chinchipé</td><td>ELISA</td><td>Serum</td><td>Cattle<br>Apparent individual prevalence</td><td>386/2668</td><td>14.5%</td><td>2668 serum samples from 386 herds</td><td>Risk factors: females (15.5 vs. 8.5%), cattle > 4 years old (20.2 vs. 10.6%)</td></tr><tr><td>Echeverría [103]</td><td>NA</td><td>2019</td><td>English</td><td>Ecuador</td><td>Quito area (Andes region) and Santo Domingo de los Tsachilas (tropical region)</td><td>ELISA</td><td>Serum</td><td>Cattle</td><td>151/352</td><td>42.9%</td><td>Dairy cattle</td><td></td></tr><tr><td>Changoluisa [104]</td><td>NA</td><td>2019</td><td>English</td><td>Ecuador</td><td>Santo Domingo Province</td><td>ELISA</td><td>Serum</td><td>Cattle<br>Cows with a first-calf abortion</td><td>91/172<br>47/95</td><td>52.9%<br>49.4%</td><td></td><td>Yes</td></tr><tr><td>Rice [105]</td><td>NA</td><td>1979</td><td>English</td><td>El Salvador</td><td>NA</td><td>CFT</td><td>Serum</td><td>Control cows<br>Cattle<br>Positive farms</td><td>44/77<br>77/297<br>27/33</td><td>57%<br>26.0%<br>82.0%</td><td></td><td>Yes</td></tr></tbody></table>
<table><caption>Table 1 (continued)</caption><thead><tr><th>Author</th><th>Year of the study</th><th>Year of publication</th><th>Language</th><th>Country</th><th>State</th><th>Methodology</th><th>Sample type</th><th>Species</th><th>Number of samples</th><th>Result</th><th>Comments</th><th>Publication in a journal</th></tr></thead><tbody><tr><td>François [106]</td><td>NA</td><td>1997</td><td>French</td><td>French Guiana</td><td>The farms tested were randomly selected on the territory</td><td>CFT</td><td></td><td>Cattle</td><td>6/355</td><td>1.7%</td><td>Representing 4% (355/9000 bovines),</td><td>Yes</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Sheep</td><td>0/171</td><td>0%</td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Goats</td><td>0/108</td><td>0%</td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>Gordon [107]</td><td>NA</td><td>2001</td><td>English</td><td>French Guiana</td><td></td><td>CFT</td><td>Serum</td><td>Cattle</td><td>6/355</td><td>1.7%</td><td></td><td>Yes</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Sheep</td><td>0/50</td><td>0%</td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Goats</td><td>0/21</td><td>0%</td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Swine</td><td>0/25</td><td>0%</td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Cattle</td><td>0/179</td><td>0%</td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Sheep</td><td>0/37</td><td>0%</td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Goats</td><td>0/16</td><td>0%</td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Swine</td><td>2/103</td><td>0 to 1.9%</td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Horses</td><td>3/88</td><td>1.1 to 3.4%</td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>Davoust [109]</td><td>2013</td><td>2014</td><td>English</td><td>French Guiana</td><td>Montagne du tigre, Cayenne</td><td>qPCR (IS1111) Serodiag-nosis</td><td>Vaginal swabs Milk Serum</td><td>Goats</td><td>0/13</td><td>0%</td><td>Doubtful serologies considered positive: 2/2 for swine; 2/3 for horses</td><td>Yes</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Sheep</td><td>0/8</td><td>0%</td><td></td><td>Investigation close to a human cluster of Q fever</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Sheep or goat</td><td>0/1</td><td>0%</td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Goats and sheep</td><td>0/37</td><td>0%</td><td></td><td>Goats and sheep are small ruminants maintained near the outbreak area</td></tr></tbody></table>

<figure><img src="image_11.png" /></figure>
<table><caption>Table 1 (continued)</caption><thead><tr><th>Author</th><th>Year of the study</th><th>Year of publication</th><th>Language</th><th>Country</th><th>State</th><th>Methodology</th><th>Sample type</th><th>Species</th><th>Number of samples</th><th>Result</th><th>Comments</th><th>Publication in a journal</th></tr></thead><tbody><tr><td>Roch</td><td>2019</td><td>2019</td><td>French</td><td>French Guiana</td><td>Montsinery</td><td>ELISA</td><td>Serum</td><td>Cattle</td><td>0/31</td><td>0%</td><td>Herd of a farmer who was infected with Q fever</td><td>No</td></tr><tr><td>Saout [110]</td><td>2015–2017</td><td>2022</td><td>English</td><td>French Guiana</td><td>Cayenne, Kourou, Sinnamary</td><td>ELISA</td><td>Serum</td><td>Cattle</td><td>152/834</td><td>14.2%</td><td></td><td>Yes</td></tr><tr><td>Johnson [101]</td><td>NA</td><td>2020</td><td>English</td><td>Grenada</td><td>NA</td><td>IFA</td><td>Serum</td><td>Cattle</td><td>0/4</td><td>0%</td><td></td><td>Yes</td></tr><tr><td>Richter and Cubes [105]</td><td>NA</td><td>1966</td><td>English</td><td>Guatemala</td><td>NA</td><td>NA</td><td>Serum</td><td>Cattle</td><td>0/86</td><td>0%</td><td></td><td></td></tr><tr><td>Salinas-Meléndez [111]</td><td>NA</td><td>2002</td><td>English</td><td>Mexico</td><td>Nuevo Leon State</td><td>ELISA</td><td>Serum</td><td>Dairy cattle</td><td>126/450</td><td>28%</td><td></td><td>Yes</td></tr><tr><td>Salman [112]</td><td>NA</td><td>1990</td><td>English</td><td>Mexico</td><td>Ensenada and Tijuana, Baja California State</td><td>ELISA</td><td>Serum</td><td>Beef cattle</td><td>19/190</td><td>10%</td><td></td><td>Cited in Rice et al. but not found</td></tr><tr><td>Johnson [101]</td><td>NA</td><td>2020</td><td>English</td><td>Montserrat</td><td>NA</td><td>IFA</td><td>Serum</td><td>Sheep</td><td>36/90</td><td>40%</td><td></td><td></td></tr><tr><td>Rojas [71]</td><td>NA</td><td>2013</td><td>Spanish</td><td>Perú</td><td>NA</td><td>RT-PCR (IS1111)</td><td>Milk</td><td>Goats</td><td>21/60</td><td>35%</td><td></td><td></td></tr><tr><td>Johnson [101]</td><td>NA</td><td>2020</td><td>English</td><td>Puerto Rico</td><td>NA</td><td>IFA</td><td>Serum</td><td>Cattle</td><td>1625/7159</td><td>22.7%</td><td></td><td></td></tr><tr><td>Johnson [101]</td><td>NA</td><td>2020</td><td>English</td><td>Saint Kitts and Nevis</td><td>St. Kitts</td><td>IFA</td><td>Serum</td><td>Positive farms</td><td>59/104</td><td>56.7%</td><td></td><td></td></tr><tr><td>Johnson [101]</td><td>NA</td><td>2020</td><td>English</td><td>Saint Kitts and Nevis</td><td>Nevis</td><td>IFA</td><td>Serum</td><td>Cattle</td><td>0/126</td><td>0%</td><td></td><td></td></tr></tbody></table>

<figure><img src="image_12.png" /></figure>
<table><caption>Table 1 (continued)</caption><thead><tr><th>Author</th><th>Year of the study</th><th>Year of publication</th><th>Language</th><th>Country</th><th>State</th><th>Methodology</th><th>Sample type</th><th>Species</th><th>Number of samples</th><th>Result</th><th>Comments</th><th>Publication in a journal</th></tr></thead><tbody><tr><td>Roch</td><td>2019</td><td>2019</td><td>French</td><td>French Guiana</td><td>Montsinery</td><td>ELISA</td><td>Serum</td><td>Cattle</td><td>0/31</td><td>0%</td><td>Herd of a farmer who was infected with Q fever</td><td>No</td></tr><tr><td>Saout [110]</td><td>2015–2017</td><td>2022</td><td>English</td><td>French Guiana</td><td>Cayenne, Kourou, Sinnamary</td><td>ELISA</td><td>Serum</td><td>Cattle</td><td>152/834</td><td>14.2%</td><td></td><td>Yes</td></tr><tr><td>Johnson [101]</td><td>NA</td><td>2020</td><td>English</td><td>Grenada</td><td>NA</td><td>IFA</td><td>Serum</td><td>Cattle</td><td>0/4</td><td>0%</td><td></td><td>Yes</td></tr><tr><td>Richter and Cubes [105]</td><td>NA</td><td>1966</td><td>English</td><td>Guatemala</td><td>NA</td><td>NA</td><td>Serum</td><td>Cattle</td><td>0/86</td><td>0%</td><td></td><td></td></tr><tr><td>Salinas-Meléndez [111]</td><td>NA</td><td>2002</td><td>English</td><td>Mexico</td><td>Nuevo Leon State</td><td>ELISA</td><td>Serum</td><td>Dairy cattle</td><td>126/450</td><td>28%</td><td></td><td>Yes</td></tr><tr><td>Salman [112]</td><td>NA</td><td>1990</td><td>English</td><td>Mexico</td><td>Ensenada and Tijuana, Baja California State</td><td>ELISA</td><td>Serum</td><td>Beef cattle</td><td>19/190</td><td>10%</td><td></td><td>Cited in Rice et al. but not found</td></tr><tr><td>Johnson [101]</td><td>NA</td><td>2020</td><td>English</td><td>Montserrat</td><td>NA</td><td>IFA</td><td>Serum</td><td>Sheep</td><td>36/90</td><td>40%</td><td></td><td></td></tr><tr><td>Rojas [71]</td><td>NA</td><td>2013</td><td>Spanish</td><td>Perú</td><td>NA</td><td>RT-PCR (IS1111)</td><td>Milk</td><td>Goats</td><td>21/60</td><td>35%</td><td></td><td></td></tr><tr><td>Johnson [101]</td><td>NA</td><td>2020</td><td>English</td><td>Puerto Rico</td><td>NA</td><td>IFA</td><td>Serum</td><td>Cattle</td><td>1625/7159</td><td>22.7%</td><td></td><td></td></tr><tr><td>Johnson [101]</td><td>NA</td><td>2020</td><td>English</td><td>Saint Kitts and Nevis</td><td>St. Kitts</td><td>IFA</td><td>Serum</td><td>Positive farms</td><td>59/104</td><td>56.7%</td><td></td><td></td></tr><tr><td>Johnson [101]</td><td>NA</td><td>2020</td><td>English</td><td>Saint Kitts and Nevis</td><td>Nevis</td><td>IFA</td><td>Serum</td><td>Cattle</td><td>0/126</td><td>0%</td><td></td><td></td></tr></tbody></table>

<figure><img src="image_11.png" /></figure>
<table><caption>**Table 1 (continued)**</caption><thead><tr><th>Author</th><th>Year of the study</th><th>Year of publication</th><th>Language</th><th>Country</th><th>State</th><th>Methodology</th><th>Sample type</th><th>Species</th><th>Number of samples</th><th>Result</th><th>Comments</th><th>Publication in a journal</th></tr></thead><tbody><tr><td>Conan [113]</td><td>2019</td><td>2020</td><td>English</td><td>Saint Kitts and Nevis</td><td>St. Kitts</td><td>ELISJA and CFT</td><td>Serum</td><td>Sheep</td><td>35/133</td><td>26.3%</td><td>Local sheep and cattle housed at a veterinary campus</td><td>Yes</td></tr><tr><td>John-son[114]</td><td>NA</td><td>2020</td><td>English</td><td>St. Lucia</td><td>NA</td><td>IFA</td><td>Serum</td><td>Cattle</td><td>0/63</td><td>0%</td><td></td><td>Yes</td></tr><tr><td>Adesiyun [114]</td><td>1992–1995</td><td>1996</td><td>English</td><td>Trinidad and Tobago</td><td>San Juan, Tunapuna and Port-of-Spain, Trinidad</td><td>CAT by Luoto</td><td>Serum</td><td>Goats</td><td>0/72</td><td>0%</td><td></td><td>Yes</td></tr><tr><td>Somma-Moreira [115, 116]</td><td>13 studies between 1956 and 1985</td><td>1987</td><td>English and Spanish</td><td>Uruguay</td><td>–</td><td>CFT, MAT</td><td>Serum</td><td>Sheep</td><td>0/18</td><td>0%</td><td>At slaughter-house</td><td>Yes</td></tr><tr><td>Macías-Rioseco [117]</td><td>2017</td><td>2019</td><td>English</td><td>Uruguay</td><td>Colonia</td><td>IHC and PCR</td><td>Placenta</td><td>Cattle</td><td>3/266</td><td>1.1%</td><td></td><td></td></tr><tr><td>Macías-Rioseco [118]</td><td>2015–2018</td><td>2019</td><td>Portuguese</td><td>Uruguay</td><td>Colonia</td><td>IHC and PCR</td><td>Fetus [53], fetus + placenta [35], and placenta only [14]</td><td>Cattle</td><td>11/256</td><td>4.3%</td><td></td><td></td></tr><tr><td>Rabaza [119]</td><td>2017</td><td>2021</td><td>English</td><td>Uruguay</td><td>San José</td><td>qPCR (1S1111), FISH</td><td>Placenta</td><td>Cattle [4]</td><td>1/1</td><td>100%</td><td>Study not on PubMed</td><td>Yes</td></tr><tr><td>Dorsch [120]</td><td>2015–2021</td><td>2022</td><td>English</td><td>Uruguay</td><td>NA</td><td>Conventional PCR</td><td>Fetus [58], fetus + placenta [36], and placenta only [6]</td><td>Sheep</td><td>0/100</td><td>0%</td><td></td><td>Yes</td></tr></tbody></table>

<figure><img src="image_13.png" /></figure>
<table><caption>Table 1 (continued)</caption><thead><tr><th>Author</th><th>Year of the study</th><th>Year of publication</th><th>Language</th><th>Country</th><th>State</th><th>Methodology</th><th>Sample type</th><th>Species</th><th>Number of samples</th><th>Result</th><th>Comments</th><th>Publication in a journal</th></tr></thead><tbody><tr><td>Conan [113]</td><td>2019</td><td>2020</td><td>English</td><td>Saint Kitts and Nevis</td><td>St. Kitts</td><td>ELISJA and CFT</td><td>Serum</td><td>Sheep</td><td>35/133</td><td>26.3%</td><td>Local sheep and cattle housed at a veterinary campus</td><td>Yes</td></tr><tr><td>John-son[114]</td><td>NA</td><td>2020</td><td>English</td><td>St. Lucia</td><td>NA</td><td>IFA</td><td>Serum</td><td>Cattle</td><td>0/63</td><td>0%</td><td></td><td>Yes</td></tr><tr><td>Adesiyun [114]</td><td>1992–1995</td><td>1996</td><td>English</td><td>Trinidad and Tobago</td><td>San Juan, Tunapuna and Port-of-Spain, Trinidad</td><td>CAT by Luoto</td><td>Serum</td><td>Goats</td><td>0/72</td><td>0%</td><td></td><td>Yes</td></tr><tr><td>Somma-Moreira [115, 116]</td><td>13 studies between 1956 and 1985</td><td>1987</td><td>English and Spanish</td><td>Uruguay</td><td>–</td><td>CFT, MAT</td><td>Serum</td><td>Sheep</td><td>0/18</td><td>0%</td><td>At slaughter-house</td><td>Yes</td></tr><tr><td>Macías-Rioseco [117]</td><td>2017</td><td>2019</td><td>English</td><td>Uruguay</td><td>Colonia</td><td>IHC and PCR</td><td>Placenta Fetuses</td><td>Cattle</td><td>3/266</td><td>1.1%</td><td></td><td></td></tr><tr><td>Macías-Rioseco [118]</td><td>2015–2018</td><td>2019</td><td>Portuguese</td><td>Uruguay</td><td>Colonia</td><td>IHC and PCR</td><td>Fetus [53], fetus + placenta [35], and placenta only [14]</td><td>Cattle</td><td>11/256</td><td>4.3%</td><td></td><td></td></tr><tr><td>Rabaza [119]</td><td>2017</td><td>2021</td><td>English</td><td>Uruguay</td><td>San José</td><td>qPCR (1S1111), FISH</td><td>Placenta Fetus</td><td>Cattle [4]</td><td>0/153</td><td>9.4%</td><td></td><td></td></tr><tr><td>Dorsch [120]</td><td>2015–2021</td><td>2022</td><td>English</td><td>Uruguay</td><td>NA</td><td>Conventional PCR</td><td>Fetus [58], fetus + placenta [36], and placenta only [6]</td><td>Sheep</td><td>5/16</td><td>0%</td><td></td><td></td></tr></tbody></table>

<figure><img src="image_12.png" /></figure>
<table><caption>**Table 1 (continued)**</caption><thead><tr><th>Author</th><th>Year of the study</th><th>Year of publication</th><th>Language</th><th>Country</th><th>State</th><th>Methodology</th><th>Sample type</th><th>Species</th><th>Number of samples</th><th>Result</th><th>Comments</th><th>Publication in a journal</th></tr></thead><tbody><tr><td>Oropeza [121]</td><td>NA</td><td>2010</td><td>Spanish</td><td>Venezuela</td><td>Villa Araure, Sabana Grande, Los Aranguez, Lara</td><td>ELISA</td><td>Serum</td><td>Goats</td><td>191/315</td><td>60.6%</td><td>Not on PubMed</td><td>Yes</td></tr></tbody></table>

A. not available; *CAT* capillary agglutination test; *CFT*, complement fixation test; *MAT*, microagglutination test; *ELISA*, enzyme-linked immunosorbent assay; *IFA*, immunofluorescence assay; *PCR*, qualitative polymerase chain reaction; *qPCR*, quantitative real-time polymerase chain reaction; *MLVA*, multilocus variable number tandem repeats analysis; *MST*, multipacer sequence type; *IHC*, immunohistochemistry; *IHA*, indirect hemagglutination; *FISH*, fluorescent in situ hybridization

goats, and sheep, were scarce. Thus, we identified one study with 25% seropositivity for *C. burnetii* in horses in Colombia in 1977 and 3.4% in French Guiana in 2007 [108, 125]. Pigs have been found to be positive in two studies: 1.9% in French Guiana in 2007, 11.3% in Trinidad in 1996, and 0 to 21.2% in Uruguay in 1984 and 1985 [115, 116], although their role in the epidemiology of *C. burnetii* remains unclear.

Finally, results vary from region to region, and it is difficult to make broad generalizations about trends on the continent. One may hypothesize a correlation between the seroprevalence of *C. burnetii* infection and animal density. The justification for the higher prevalence of cattle would be in places where there is intensive animal farming [126, 127]. In any case, estimated seroprevalence data are very approximate, which makes it necessary to work on protocols and monitoring devices for *C. burnetii* infections.

## Pets

A total of 18 studies from seven countries were performed on detecting *C. burnetii* in pets-dogs and cats -mainly in French Guiana and Brazil (Table 2). These studies included five unpublished works, one thesis in French Guiana and Brazil, respectively, and three unpublished works originating from the investigation of the epidemiology and veterinary department of the French Army following outbreaks in military personnel (Fig. 3). Dogs were found positive in the indirect immunofluorescence assay (IFA) for *C. burnetii* in Argentina, Brazil, and French Guiana, with prevalence from 1.8 to 21.7% according to the region, the study, and the cutoff of the titer of IgG [107, 128, 129, 131, 134]. In Brazil and French Guiana, some dogs have been found to be positive, with higher prevalence rates around human Q fever cases [76, 77, 107, 130]. Although some of these dogs were inquired about during confirmed human Q fever outbreak investigations, they do not seem to be at the origin of these human clusters [76, 107, 130]. Cats have been studied in only three regions: Argentina, Brazil, and French Guiana [77, 107–109, 128–131, 134] with six cats positive, five in Argentina and one in Rio de Janeiro, Brazil [77, 128]. No evidence of *C. burnetii* infection was found in dogs using serology or PCR tests on vaginal swabs in Colombia, Martinique, Nicaragua, and Uruguay [97, 134, 135].

Aside from livestock, pet animals, especially dogs and cats, in close contact with humans are important potential reservoirs of *C. burnetii* during urban Q fever outbreaks [124]. Some cases of human coxiellosis have been reported from infected dogs and cats as a source of infection [14, 138]. At the same time, several pets were found to be positive in the investigation of human clusters without these animals being incriminated in the transmission to humans [76, 107, 130].
<table><caption>**Table 1 (continued)**</caption><thead><tr><th>Author</th><th>Year of the study</th><th>Year of publication</th><th>Language</th><th>Country</th><th>State</th><th>Methodology</th><th>Sample type</th><th>Species</th><th>Number of samples</th><th>Result</th><th>Comments</th><th>Publication in a journal</th></tr></thead><tbody><tr><td>Oropeza [121]</td><td>NA</td><td>2010</td><td>Spanish</td><td>Venezuela</td><td>Villa Araure, Sabana Grande, Los Aranguez, Lara</td><td>ELISA</td><td>Serum</td><td>Goats</td><td>191/315</td><td>60.6%</td><td>Not on PubMed</td><td>Yes</td></tr></tbody></table>

A. not available; *CAT* capillary agglutination test; *CFT*, complement fixation test; *MAT*, microagglutination test; *ELISA*, enzyme-linked immunosorbent assay; *IFA*, immunofluorescence assay; *PCR*, qualitative polymerase chain reaction; *qPCR*, quantitative real-time polymerase chain reaction; *MLVA*, multilocus variable number tandem repeats analysis; *MST*, multipacer sequence type; *IHC*, immunohistochemistry; *IHA*, indirect hemagglutination; *FISH*, fluorescent in situ hybridization

goats, and sheep, were scarce. Thus, we identified one study with 25% seropositivity for *C. burnetii* in horses in Colombia in 1977 and 3.4% in French Guiana in 2007 [108, 125]. Pigs have been found to be positive in two studies: 1.9% in French Guiana in 2007, 11.3% in Trinidad in 1996, and 0 to 21.2% in Uruguay in 1984 and 1985 [115, 116], although their role in the epidemiology of *C. burnetii* remains unclear.

Finally, results vary from region to region, and it is difficult to make broad generalizations about trends on the continent. One may hypothesize a correlation between the seroprevalence of *C. burnetii* infection and animal density. The justification for the higher prevalence of cattle would be in places where there is intensive animal farming [126, 127]. In any case, estimated seroprevalence data are very approximate, which makes it necessary to work on protocols and monitoring devices for *C. burnetii* infections.

## Pets

A total of 18 studies from seven countries were performed on detecting *C. burnetii* in pets-dogs and cats -mainly in French Guiana and Brazil (Table 2). These studies included five unpublished works, one thesis in French Guiana and Brazil, respectively, and three unpublished works originating from the investigation of the epidemiology and veterinary department of the French Army following outbreaks in military personnel (Fig. 3). Dogs were found positive in the indirect immunofluorescence assay (IFA) for *C. burnetii* in Argentina, Brazil, and French Guiana, with prevalence from 1.8 to 21.7% according to the region, the study, and the cutoff of the titer of IgG [107, 128, 129, 131, 134]. In Brazil and French Guiana, some dogs have been found to be positive, with higher prevalence rates around human Q fever cases [76, 77, 107, 130]. Although some of these dogs were inquired about during confirmed human Q fever outbreak investigations, they do not seem to be at the origin of these human clusters [76, 107, 130]. Cats have been studied in only three regions: Argentina, Brazil, and French Guiana [77, 107–109, 128–131, 134] with six cats positive, five in Argentina and one in Rio de Janeiro, Brazil [77, 128]. No evidence of *C. burnetii* infection was found in dogs using serology or PCR tests on vaginal swabs in Colombia, Martinique, Nicaragua, and Uruguay [97, 134, 135].

Aside from livestock, pet animals, especially dogs and cats, in close contact with humans are important potential reservoirs of *C. burnetii* during urban Q fever outbreaks [124]. Some cases of human coxiellosis have been reported from infected dogs and cats as a source of infection [14, 138]. At the same time, several pets were found to be positive in the investigation of human clusters without these animals being incriminated in the transmission to humans [76, 107, 130].
## Wildlife

A total of 24 studies, to which we must add the unpublished results of two investigations, were published on wildlife from only five countries of Latin America: Chile, Colombia, Brazil, French Guiana, and Uruguay (Fig. 4). Some studies have been performed through serodiagnosis in various mammal species, and most of them were negative, as shown in Table 3. Nevertheless, some species have shown seropositivity, meaning contact with the bacterium, such as free-living cervids (*Blastocerus dichotomous* and *Mazama gouazoubira*) in Mato Grasso do Sul, São Paulo, Goiás, and Paraná, Brazil, rodents and marsupials in French Guiana (*Proechimys* sp., *Philander opossum*, *Didelphis marsupialis* [107, 151]) and Pampas deer (*Ozotocero sbezoarticus*) in the Department of Maldonado of Uruguay. Thus, recent studies conducted in Brazil showed serological evidence of *C. burnetii* infection in *Wiedomys pyrrhorhinos* rodents and *Didelphis albiventris* marsupial from Ceará and Pernambuco states, respectively [131]. These animals were collected from areas where dogs were also found seroreactive to *C. burnetii*, suggesting that wild and peridomestic cycles of *C. burnetii* can be connected by rodents and other wild mammals, a scenario identified in other regions where peridomestic rather than wild cycles have a high impact in coxiellosis cases [131, 157]. The use of molecular biology, mainly PCR targeting the repetitive element *IS1111*, helped identify new positive mammals (Table 4). The results reported in Ct (cycle threshold) were mostly superior to 35, which corresponds to traces in bacterial loads. The animal groups with molecular evidence of *C. burnetii* in Latin America were Chiroptera (7 species), Rodentia (6 species), Suina (2 species), and one species for Pilosa, Cingulata, and Perissodactyla, respectively (Table 4).

**Nonflying Mammals** In French Guiana, the first studies about *C. burnetii* infection in wildlife date back to the late 1990s, following the discovery of the first human cases. This enthusiasm for the search for an animal reservoir of *C. burnetii* in this small French territory of about 300,000 inhabitants (200,000 inhabitants at the time of the first investigations) and an area of 83,846 km² is due to the real public health burden represented by human Q fever and the occurrence of several clusters in the French military personnel, linked to a very proactive Nation Reference Center in Marseille, in mainland France. Furthermore, these investigations have been enhanced, as all the first studies on animal infection with *C. burnetii* seem to rule out cattle as a source of human infection, so the origin of the epidemics had to be found in wildlife [107, 108, 158]. Although a positive three-toed sloth (*Bradypus tridactylus*) was found positive [109, 154], other animals, such as capybara (*Hydrochoerus hydrochaeris*), were tested and found negative [109, 152], weakening the former hypothesis claiming that the sloth was the single

reservoir [151, 159]. Indeed, other species have been found positive between 2005 and 2014 following numerous investigations [153], and several species were found to be positive through the analysis by *IS1111* PCR of the muscular juice of animals collected from hunters. Thus, one nine-banded armadillo (*Dasypus novemcinctus*), three white-lipped peccaries (*Tayassu pecari*), three collared peccaries (*Pecari tajacu*), one south American tapir (*Tapirus terrestris*), and two other capybaras (*Hydrochoerus hydrochaeris*) have already been found to be positive, most of them with Ct superior to 35, but never published. One spiny rat (*Proechimys cuvieri*) was also found positive on liver samples among 29 other rodents [153]. A study is currently underway in French Guiana that has collected over 1000 samples from animals, especially tissues and stools, to better characterize the wildlife reservoir of *C. burnetii* [160].

In Brazil, the first study found on the search for *C. burnetii* in wildlife is a thesis, which is an investigation in domestic animals, wild animals, and arthropods in Itaboraí in the state of Rio de Janeiro around cases of suspected *C. burnetii* infection [77]. Afterward, in a study developed in eight municipalities in the state of Rio de Janeiro, in the Atlantic Forest, between 2007 and 2012, the prevalence of *C. burnetii* DNA in 131 different species of wild rodents was 4.6% [139]. The positive rodents belonged to the species *Akodon cursor* (3/32: 9.4%), *Mus musculus* (1/19: 5.3%), *Oxymycterus dasytrichus* (1/12: 8.3%), and *Oligoryzomys nigripes* (1/16: 6.3%) captured in forest edges and near human dwellings in the municipalities of Piraí and Valença [139]. Other studies have been conducted on other species of nonflying mammals, but all with negative results. Thus studies using PCR have also been performed in several taxa: free-living cervids, free-living wild boars, and different species of Xenarthra in Brazil [141, 145, 146], Darwin's fox (*Lycalopex fulvipes*) in Chile [149], different species of wild canids, rodents, and marsupials in Brazil, French Guiana, and Uruguay [107, 109, 115, 116, 131, 142], and in reptiles and amphibians in French Guiana without the detection of *C. burnetii* DNA [107, 109, 143].

**Chiroptera** Concerning bats, although the role of these animals in the enzootic cycles of *C. burnetii* has received little attention worldwide, it was possible to find studies developed in Brazil and Chile [147•]. The first study to characterize this proteobacteria in Latin America involved 119 bats from 21 species captured in preserved areas in Rio de Janeiro, Bahia, and Santa Catarina, Brazilian from 2014 to 2015. *Coxiella burnetii* was PCR-detected in specimens of two species from the genus *Artibeus*: the great fruit-eating bat (*Artibeus lituratus*) (3 *C. burnetii*-positive/14 bats) and the fringed fruit-eating bat (*Artibeus fimbriatus*) (1/7) from two different regions: Jacarepaguá (Rio de Janeiro state) (3/44; 7%) and Serra do Tabuleiro State Park (Santa Catarina state) (1/28; 4%) [140]. In the second study, the first report of *C. burnetii* in Chile, PCR positivity of 9.0% (5/55) was observed in Mexican free-tailed bats or Brazilian free-tailed bats (*Tadarida brasiliensis*) sampled in three regions of

<figure><img src="image_14.png" /></figure>
Wildlife

A total of 24 studies, to which we must add the unpublished
results of two investigations, were published on wildlife
from only five countries of Latin America: Chile, Colom-
bia, Brazil, French Guiana, and Uruguay (Fig. 4). Some
studies have been performed through serodiagnosis in vari-
ous mammal species, and most of them were negative, as
shown in Table 3. Nevertheless, some species have shown
seropositivity, meaning contact with the bacterium, such as
free-living cervids (Blastocerus dichotomous and Mazama
gouazoubira) in Mato Grasso do Sul, São Paulo, Goiás, and
Paraná, Brazil, rodents and marsupials in French Guiana
(Proechimys sp., Philander opossum, Didelphis marsupia-
lis [107, 151]) and Pampas deer (Ozotocero sbezoarticus)
in the Department of Maldonado of Uruguay. Thus, recent
studies conducted in Brazil showed serological evidence of
C. burnetii infection in Wiedomys pyrrhorhinos rodents and
Didelphis albiventris marsupial from Ceará and Pernambuco
states, respectively [131]. These animals were collected from
areas where dogs were also found seroreactive to C. burnetii,
suggesting that wild and peridomestic cycles of C. burnetii
can be connected by rodents and other wild mammals, a sce-
nario identified in other regions where peridomestic rather
than wild cycles have a high impact in coxiellosis cases [131,
157]. The use of molecular biology, mainly PCR targeting
the repetitive element IS1111, helped identify new positive
mammals (Table 4). The results reported in Ct (cycle tre-
shold) were mostly superior to 35, which corresponds to
traces in bacterial loads. The animal groups with molecular
evidence of C. burnetii in Latin America were Chiroptera
(7 species), Rodentia (6 species), Suina (2 species), and one
species for Pilosa, Cingulata, and Perissodactyla, respec-
tively (Table 4).

**Nonflying Mammals** In French Guiana, the first studies about *C. burnetii* infection in wildlife date back to the late 1990s, following the discovery of the first human cases. This enthusiasm for the search for an animal reservoir of *C. burnetii* in this small French territory of about 300,000 inhabitants (200,000 inhabitants at the time of the first investigations) and an area of 83,846 km² is due to the real public health burden represented by human Q fever and the occurrence of several clusters in the French military personnel, linked to a very proactive Nation Reference Center in Marseille, in mainland France. Furthermore, these investigations have been enhanced, as all the first studies on animal infection with *C. burnetii* seem to rule out cattle as a source of human infection, so the origin of the epidemics had to be found in wildlife [107, 108, 158]. Although a positive three-toed sloth (*Bradypus tridactylus*) was found positive [109, 154], other animals, such as capybara (*Hydrochoerus hydrochaeris*), were tested and found negative [109, 152], weakening the former hypothesis claiming that the sloth was the single

reservoir [151, 159]. Indeed, other species have been found posi-
tive between 2005 and 2014 following numerous investigations
[153], and several species were found to be positive through the
analysis by IS1111 PCR of the muscular juice of animals col-
lected from hunters. Thus, one nine-banded armadillo (Dasypus
novemcinctus), three white-lipped peccaries (Tayassu pecari),
three collared peccaries (Pecari tajacu), one south American
tapir (Tapirus terrestris), and two other capybaras (Hydrochoe-
rus hydrochaeris) have already been found to be positive, most
of them with Ct superior to 35, but never published. One spiny
rat (Proechimys cuvieri) was also found positive on liver samples
among 29 other rodents [153]. A study is currently underway
in French Guiana that has collected over 1000 samples from
animals, especially tissues and stools, to better characterize the
wildlife reservoir of C. burnetii [160].

In Brazil, the first study found on the search for *C. burnetii*
in wildlife is a thesis, which is an investigation in domestic ani-
mals, wild animals, and arthropods in Itaboraí in the state of Rio
de Janeiro around cases of suspected *C. burnetii* infection [77].
Afterward, in a study developed in eight municipalities in the
state of Rio de Janeiro, in the Atlantic Forest, between 2007 and
2012, the prevalence of *C. burnetii* DNA in 131 different species
of wild rodents was 4.6% [139]. The positive rodents belonged
to the species *Akodon cursor* (3/32: 9.4%), *Mus musculus* (1/19:
5.3%), *Oxymycterus dasytrichus* (1/12: 8.3%), and *Oligoryzomys
nigripes* (1/16: 6.3%) captured in forest edges and near human
dwellings in the municipalities of Piraí and Valença [139]. Other
studies have been conducted on other species of nonflying mam-
mals, but all with negative results. Thus studies using PCR have
also been performed in several taxa: free-living cervids, free-
living wild boars, and different species of Xenarthra in Brazil
[141, 145, 146], Darwin’s fox (*Lycalopex fulvipes*) in Chile
[149], different species of wild canids, rodents, and marsupials
in Brazil, French Guiana, and Uruguay [107, 109, 115, 116, 131,
142], and in reptiles and amphibians in French Guiana without
the detection of *C. burnetii* DNA [107, 109, 143].

**Chiroptera** Concerning bats, although the role of these animals in the enzootic cycles of *C. burnetii* has received little attention worldwide, it was possible to find studies developed in Brazil and Chile [147•]. The first study to characterize this proteobacteria in Latin America involved 119 bats from 21 species captured in preserved areas in Rio de Janeiro, Bahia, and Santa Catarina, Brazilian from 2014 to 2015. *Coxiella burnetii* was PCR-detected in specimens of two species from the genus *Artibeus*: the great fruit-eating bat (*Artibeus lituratus*) (3 *C. burnetii*-positive/14 bats) and the fringed fruit-eating bat (*Artibeus fimbriatus*) (1/7) from two different regions: Jacarepaguá (Rio de Janeiro state) (3/44; 7%) and Serra do Tabuleiro State Park (Santa Catarina state) (1/28; 4%) [140]. In the second study, the first report of *C. burnetii* in Chile, PCR positivity of 9.0% (5/55) was observed in Mexican free-tailed bats or Brazilian free-tailed bats (*Tadarida brasiliensis*) sampled in three regions of

<figure><img src="image_13.png" /></figure>
the country [147•]. Lastly, a third study was published in 2022 using *IS1111*-qPCR to detect *C. burnetii* in blood samples from 126 bats captured in 2014, 2015, and 2018 in the Macaregua cave, Colombia [161]. Molecular evidence of *C. burnetii* was found in 6.3% of the samples: 3/49 (6.1%) Seba's short-tailed bats (*Carollia perspicillata*), a widespread frugivorous bat; 2/35 (5.7%) ghost-faced bats (*Mormoops megalophylla*); and 3/42 (7.1%) Trinidadian funnel-eared bats (*Natalus tumidirostris*), an insectivorous bat.

There is a strong popular rumor that bats are reservoirs and transmitters of *C. burnetii* to humans in French Guiana. However, no study supported this hypothesis, and it is difficult to know where this rumor originated. In one of the very first studies conducted in the late 1990s, it was shown that seeing bats near one's home was independently associated with the occurrence of Q fever in patients with fever compared to patients supported for dengue fever [107]. Nevertheless, the same team that found the various nonflying mammals positive found six bats belonging to four species positive to *C. burnetii*, spread over the whole territory but with high Ct (> 35): three Seba's short-tailed bat (*Carollia perspicillata*), one Parnell's mustached bat (*Pteronotus parnellii*), a lesser spear-nosed bat (*Phyllostomus elongatus*), and a lesser bulldog bat (*Noctilio albiventris*) [153].

Thus, even though the potential role of these animals as a source of infection for humans and other animals, including cattle, is still unknown, these reports suggest the existence of a complex *C. burnetii* transmission cycle involving a large number of wild mammals [162]. Therefore, further studies must be conducted to better understand their role in *C. burnetii* cycles. The results of this research on species other than domestic ruminants are difficult to interpret because the choices and times of the samples are not homogeneous, and the PCR tests performed in this way on a wide variety of biological matrices (various organs and fluids) allow first-line screening but not a controlled quantification for analytical epidemiology. Methodological developments are required.

## Ticks and Ectoparasites

There has been an increasing interest in tick-borne pathogens, including *C. burnetii*, these last few years (Fig. 5). Eleven countries have published at least one study about *C. burnetii* in ticks in Latin America and the Caribbean, with a huge number of studies coming from Brazil, especially in the last 3 years (N=19), followed by French Guiana (N=7) and Colombia (N=5). Although the importance of ticks in the epidemiology of *C. burnetii* infection remains debatable [9], there is no doubt that ticks can be infected by *C. burnetii* and that they can, therefore, act as vectors [6, 163–165]. In natural conditions, several *C. burnetii* strains have been successfully isolated from wild ticks, including from a few

South American tick species (Table 5) [6, 173]. The Fiocruz group published a study in which *Amblyomma sculptum* and *Rhipicephalus sanguineus* were PCR-positive for *C. burnetii* [173]. For instance, in the case study by Pacheco et al. (2013) in Argentina, *C. burnetii* infection in two tick species belonging to the *Amblyomma* genus, *A. tigrinum* and *A. parvum*, was confirmed using hemolymph tests, isolation in Vero cells, and multilocus DNA sequencing. The strain At12 was typed as ST 73 in this publication. In Cuba, Noda and colleagues found six *Amblyomma mixtum* from 2 tick pools positive for *C. burnetii* using IS1111 qPCR [184]. In Brazil and French Guiana, specimens of another *Amblyomma* tick species, *A. geayi*, collected on three-toed sloth infected by *C. burnetii* were also infected [109, 167]. Therefore, a sylvatic cycle based on *C. burnetii* tick-borne transmission seems to be sustainable. However, *C. burnetii* is probably far more frequently transmitted through the airborne route than through ticks [6]. Although most of the studies did not find any sign of infection by *C. burnetii* in South American tick species, recent observations, based on advances in molecular and cell biology, showed that ticks, including those found in South America, commonly harbor *Coxiella*-like endosymbionts (*Coxiella*-LE), closely related but genetically distinct to *C. burnetii*. These *Coxiella*-LE are almost exclusively confined to ticks and, according to current knowledge, pose a much lower infection risk to vertebrates than *C. burnetii* [6]. Extensive molecular surveys have consistently revealed that *Coxiella*-LE predominates in most tick species investigated thus far, with at least two-thirds of tick species being naturally infected [156, 168, 187, 191]. Most importantly, *Coxiella*-LE has been commonly misidentified as *C. burnetii* [6, 192••, 193, 194]. Several *C. burnetii* detection methods are in use, but many are not efficient enough to clearly distinguish between *C. burnetii* and *Coxiella*-LE [192••, 193–195]. Based on the identification of IS1111 in both *C. burnetii* and the *Coxiella*-LE, Mares-Guia et al. (2018) developed a nested PCR assay, considering that IS1111 amino-acid sequences in endosymbionts revealed to be genetically divergent, including degraded copies likely to be nonfunctional, providing a specific *C. burnetii* detection [173, 194].

## Highlights on the Epidemiological Situation of Specifically Selected Countries

### French Guiana

French Guiana is a French overseas territory located in the northeast corner of the South American continent. The territory of 83,534 km² is more than 90% covered by the Amazon rainforest, and the population of approximately 300,000 inhabitants (www.insee.fr) is concentrated on the coast, particularly in Cayenne and its surroundings. The human
the country [147]. Lastly, a third study was published in 2022 using *IS1111*-qPCR to detect *C. burnetii* in blood samples from 126 bats captured in 2014, 2015, and 2018 in the Macaregua cave, Colombia [161]. Molecular evidence of *C. burnetii* was found in 6.3% of the samples: 3/49 (6.1%) Seba's short-tailed bats (*Carollia perspicillata*), a widespread frugivorous bat; 2/35 (5.7%) ghost-faced bats (*Mormoops megalophylla*); and 3/42 (7.1%) Trinidadian funnel-eared bats (*Natalus tumidirostris*), an insectivorous bat.

There is a strong popular rumor that bats are reservoirs and transmitters of *C. burnetii* to humans in French Guiana. However, no study supported this hypothesis, and it is difficult to know where this rumor originated. In one of the very first studies conducted in the late 1990s, it was shown that seeing bats near one's home was independently associated with the occurrence of Q fever in patients with fever compared to patients supported for dengue fever [107]. Nevertheless, the same team that found the various nonflying mammals positive found six bats belonging to four species positive to *C. burnetii*, spread over the whole territory but with high Ct (> 35): three Seba's short-tailed bat (*Carollia perspicillata*), one Parnell's mustached bat (*Pteronotus parnellii*), a lesser spear-nosed bat (*Phyllostomus elongatus*), and a lesser bulldog bat (*Noctilio albiventris*) [153].

Thus, even though the potential role of these animals as a source of infection for humans and other animals, including cattle, is still unknown, these reports suggest the existence of a complex *C. burnetii* transmission cycle involving a large number of wild mammals [162]. Therefore, further studies must be conducted to better understand their role in *C. burnetii* cycles. The results of this research on species other than domestic ruminants are difficult to interpret because the choices and times of the samples are not homogeneous, and the PCR tests performed in this way on a wide variety of biological matrices (various organs and fluids) allow first-line screening but not a controlled quantification for analytical epidemiology. Methodological developments are required.

## Ticks and Ectoparasites

There has been an increasing interest in tick-borne pathogens, including *C. burnetii*, these last few years (Fig. 5). Eleven countries have published at least one study about *C. burnetii* in ticks in Latin America and the Caribbean, with a huge number of studies coming from Brazil, especially in the last 3 years (N=19), followed by French Guiana (N=7) and Colombia (N=5). Although the importance of ticks in the epidemiology of *C. burnetii* infection remains debatable [9], there is no doubt that ticks can be infected by *C. burnetii* and that they can, therefore, act as vectors [6, 163–165]. In natural conditions, several *C. burnetii* strains have been successfully isolated from wild ticks, including from a few

South American tick species (Table 5) [6, 173]. The Fiocruz group published a study in which *Amblyomma sculptum* and *Rhipicephalus sanguineus* were PCR-positive for *C. burnetii* [173]. For instance, in the case study by Pacheco et al. (2013) in Argentina, *C. burnetii* infection in two tick species belonging to the *Amblyomma* genus, *A. tigrinum* and *A. parvum*, was confirmed using hemolymph tests, isolation in Vero cells, and multilocus DNA sequencing. The strain At12 was typed as ST 73 in this publication. In Cuba, Noda and colleagues found six *Amblyomma mixtum* from 2 tick pools positive for *C. burnetii* using *IS1111* qPCR [184]. In Brazil and French Guiana, specimens of another *Amblyomma* tick species, *A. geayi*, collected on three-toed sloth infected by *C. burnetii* were also infected [109, 167]. Therefore, a sylvatic cycle based on *C. burnetii* tick-borne transmission seems to be sustainable. However, *C. burnetii* is probably far more frequently transmitted through the airborne route than through ticks [6]. Although most of the studies did not find any sign of infection by *C. burnetii* in South American tick species, recent observations, based on advances in molecular and cell biology, showed that ticks, including those found in South America, commonly harbor *Coxiella*-like endosymbionts (*Coxiella*-LE), closely related but genetically distinct to *C. burnetii*. These *Coxiella*-LE are almost exclusively confined to ticks and, according to current knowledge, pose a much lower infection risk to vertebrates than *C. burnetii* [6]. Extensive molecular surveys have consistently revealed that *Coxiella*-LE predominates in most tick species investigated thus far, with at least two-thirds of tick species being naturally infected [156, 168, 187, 191]. Most importantly, *Coxiella*-LE has been commonly misidentified as *C. burnetii* [6, 192••, 193, 194]. Several *C. burnetii* detection methods are in use, but many are not efficient enough to clearly distinguish between *C. burnetii* and *Coxiella*-LE [192••, 193–195]. Based on the identification of *IS1111* in both *C. burnetii* and the *Coxiella*-LE, Mares-Guia et al. (2018) developed a nested PCR assay, considering that *IS1111* amino-acid sequences in endosymbionts revealed to be genetically divergent, including degraded copies likely to be nonfunctional, providing a specific *C. burnetii* detection [173, 194].

## Highlights on the Epidemiological Situation of Specifically Selected Countries

### French Guiana

French Guiana is a French overseas territory located in the northeast corner of the South American continent. The territory of 83,534 km² is more than 90% covered by the Amazon rainforest, and the population of approximately 300,000 inhabitants (www.insee.fr) is concentrated on the coast, particularly in Cayenne and its surroundings. The human
<figure><img src="image_15.png" /><figcaption>**Table 2** Data on *Coxiella burnetii* in pets in Latin America</figcaption></figure>

<table><thead><tr><th>Author</th><th>Year of the study</th><th>Year of publication</th><th>Language</th><th>Country</th><th>State</th><th>Biological methodology</th><th>Sample type</th><th>Species</th><th>Number of samples</th><th>Result</th><th>Comments</th><th>Published</th></tr></thead><tbody><tr><td>Navarro O'Connor [128]</td><td>2001–2004</td><td>2007</td><td>Spanish</td><td>Argentina</td><td>Buenos Aires</td><td>IFA</td><td>Serum</td><td>Cats</td><td>1–5/55</td><td>1.8–9%</td><td>1 to 5 positive according to the titer used for IgG</td><td>Yes</td></tr><tr><td>Cicuttin [129]</td><td>2002–2003</td><td>2013</td><td>Spanish</td><td>Argentina</td><td>Buenos Aires</td><td>IFA</td><td>Serum</td><td>Dogs</td><td>0/16</td><td>0%</td><td>Yes</td><td>Yes</td></tr><tr><td>Lemos [130]</td><td>2008</td><td>2011</td><td>English</td><td>Brazil</td><td>Rio de Janeiro</td><td>IFA</td><td>Serum</td><td>Dogs</td><td>3–19/123</td><td>2.4–15.4%</td><td>15.4% (cut-off 1.50)</td><td>Yes</td></tr><tr><td>Mares-Guia [76]</td><td>2008–2009</td><td>2014</td><td>English</td><td>Brazil</td><td>City of Itaboraí, Rio de Janeiro State</td><td>IFA</td><td>Serum</td><td>Dogs</td><td>2/14</td><td>14.2%</td><td>Area of the first human case of Q fever in Brazil</td><td>Yes</td></tr><tr><td>Mares-Guia [77]</td><td>2011–2012</td><td>2015</td><td>Portuguese</td><td>Brazil</td><td>Rio de Janeiro</td><td>IFA</td><td>Serum</td><td>Cats</td><td>0/1</td><td>0%</td><td>Yes</td><td>No</td></tr><tr><td>Barboza de Oliveira [131]</td><td>2014–2016</td><td>2020</td><td>English</td><td>Brazil</td><td>Ceara and Pernambuco (Caatinga biome)</td><td>IFA</td><td>Anal swab</td><td>Cats</td><td>0/1</td><td>0%</td><td>Thesis. Unpublished results</td><td>Yes</td></tr><tr><td>Oliveira [132]</td><td>2016–2019</td><td>2021</td><td>English</td><td>Brazil</td><td>Town of Barra Mansa, Rio de Janeiro State</td><td>PCR (hpAB)</td><td>Serum</td><td>Dogs</td><td>3/13</td><td>23.0%</td><td>Yes</td><td>Yes</td></tr><tr><td>Di Cataldo [133]</td><td>2006–2019</td><td>2022</td><td>English</td><td>Chile</td><td>4 different areas: coastal desert, mountain desert, Steppe-Mediterranean, and Temperate Warm Rainy</td><td>IFA</td><td>Serum</td><td>Dogs</td><td>5/147</td><td>3.4%</td><td>Dogs positive in PCR and IgG positive for L. infantum</td><td>Yes</td></tr><tr><td>Faccini-Martínez [97]</td><td>2011</td><td>2017</td><td>English</td><td>Colombia</td><td>Villeta, Cundinamarca</td><td>IFA</td><td>Serum</td><td>Dogs</td><td>0/118</td><td>0%</td><td>Yes</td><td>Yes</td></tr><tr><td>Boni [134]</td><td>NA</td><td>1998</td><td>English</td><td>French Guiana</td><td>NA</td><td>IFA</td><td>Serum</td><td>Dogs</td><td>1/19</td><td>5.2%</td><td>Dogs belonging to the French military</td><td>Yes</td></tr><tr><td>Gardon [107]</td><td>NA</td><td>2001</td><td>English</td><td>French Guiana</td><td>Cayenne area</td><td>IFA</td><td>Serum</td><td>Dogs</td><td>7/57</td><td>12.3%</td><td>25/57 and 6/6 belonging to humans with proven Q fever, respectively</td><td>Yes</td></tr></tbody></table>
<figure><img src="image_14.png" /></figure>

<table><caption>**Table 2** Data on *Coxiella burnetii* in pets in Latin America</caption><thead><tr><th>Author</th><th>Year of the study</th><th>Year of publication</th><th>Language</th><th>Country</th><th>State</th><th>Biological methodology</th><th>Sample type</th><th>Species</th><th>Number of samples</th><th>Result</th><th>Comments</th><th>Published</th></tr></thead><tbody><tr><td>Navarro O'Connor [128]</td><td>2001–2004</td><td>2007</td><td>Spanish</td><td>Argentina</td><td>Buenos Aires</td><td>IFA</td><td>Serum</td><td>Cats</td><td>1–5/55</td><td>1.8–9%</td><td>1 to 5 positive according to the titer used for IgG</td><td>Yes</td></tr><tr><td>Cicuttin [129]</td><td>2002–2003</td><td>2013</td><td>Spanish</td><td>Argentina</td><td>Buenos Aires</td><td>IFA</td><td>Serum</td><td>Dogs</td><td>0/16</td><td>0%</td><td>Yes</td><td>Yes</td></tr><tr><td>Lemos [130]</td><td>2008</td><td>2011</td><td>English</td><td>Brazil</td><td>Rio de Janeiro</td><td>IFA</td><td>Serum</td><td>Dogs</td><td>3–19/123</td><td>2.4–15.4%</td><td>15.4% (cut-off 1.50)</td><td>Yes</td></tr><tr><td>Mares-Guia [76]</td><td>2008–2009</td><td>2014</td><td>English</td><td>Brazil</td><td>City of Itaboraí, Rio de Janeiro State</td><td>IFA</td><td>Serum</td><td>Dogs</td><td>2/14</td><td>14.2%</td><td>Area of the first human case of Q fever in Brazil</td><td>Yes</td></tr><tr><td>Mares-Guia [77]</td><td>2011–2012</td><td>2015</td><td>Portuguese</td><td>Brazil</td><td>Rio de Janeiro</td><td>IFA</td><td>Serum</td><td>Cats</td><td>0/1</td><td>0%</td><td>Yes</td><td>No</td></tr><tr><td>Barboza de Oliveira [131]</td><td>2014–2016</td><td>2020</td><td>English</td><td>Brazil</td><td>Ceara and Pernambuco (Caatinga biome)</td><td>IFA</td><td>Serum</td><td>Cats</td><td>0/1</td><td>0%</td><td>Yes</td><td>Yes</td></tr><tr><td>Oliveira [132]</td><td>2016–2019</td><td>2021</td><td>English</td><td>Brazil</td><td>Town of Barra Mansa, Rio de Janeiro State</td><td>IFA</td><td>Serum</td><td>Cats</td><td>3/13</td><td>23.0%</td><td>Thesis. Unpublished results</td><td>No</td></tr><tr><td>Di Cataldo [133]</td><td>2006–2019</td><td>2022</td><td>English</td><td>Chile</td><td>4 different areas: coastal desert, mountain desert, Steppe-Mediterranean, and Temperate Warm Rainy</td><td>IFA</td><td>Serum</td><td>Dogs</td><td>5/147</td><td>3.4%</td><td>Yes</td><td>Yes</td></tr><tr><td>Faccini-Martínez [97]</td><td>2011</td><td>2017</td><td>English</td><td>Colombia</td><td>Villeta, Cundinamarca</td><td>IFA</td><td>Serum</td><td>Dogs</td><td>0/358</td><td>0%</td><td>Dogs positive in PCR and IgG positive for L. infantum</td><td>Yes</td></tr><tr><td>Boni [134]</td><td>NA</td><td>1998</td><td>English</td><td>French Guiana</td><td>NA</td><td>IFA</td><td>Serum</td><td>Dogs</td><td>0/118</td><td>0%</td><td>Yes</td><td>Yes</td></tr><tr><td>Gardon [107]</td><td>NA</td><td>2001</td><td>English</td><td>French Guiana</td><td>Cayenne area</td><td>IFA</td><td>Serum</td><td>Dogs</td><td>7/57</td><td>12.3%</td><td>Dogs belonging to the French military</td><td>Yes</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Cats</td><td>0/6</td><td>0%</td><td>Yes</td><td>Yes</td></tr></tbody></table>
<table><caption>**Table 2 (continued)**</caption><thead><tr><th>Author</th><th>Year of the study</th><th>Year of publication</th><th>Language</th><th>Country</th><th>State</th><th>Biological methodology</th><th>Sample type</th><th>Species</th><th>Number of samples</th><th>Result</th><th>Comments</th><th>Published</th></tr></thead><tbody><tr><td>Debin [108]</td><td>2007</td><td>2007</td><td>French</td><td>French Guiana</td><td>Communes du littoral</td><td>ELISA</td><td>Serum</td><td>Dogs</td><td>16/63 (including 8 doubtful cases, 4 positive animals, and 2 positive animals belonging to the same owner</td><td>9.2 to 21.7%</td><td>22 from Kourou kennels and 41 from veterinarians</td><td>Yes</td></tr><tr><td>Davoust</td><td>2013</td><td>2014</td><td>French</td><td>French Guiana</td><td>Montagne du Tigre, Cayenne</td><td>qPCR (IS1111)</td><td>Vaginal swabs</td><td>Dogs</td><td>0/3</td><td>0%</td><td>Unpublished data</td><td>No</td></tr><tr><td>Davoust</td><td>2013</td><td>2014</td><td>French</td><td>French Guiana</td><td>IFA</td><td>IFA</td><td>Serum</td><td>Cat</td><td>0/1</td><td>0%</td><td></td><td></td></tr><tr><td>Davoust [109]</td><td>2013</td><td>2014</td><td>English</td><td>French Guiana</td><td>Cayenne and Kourou</td><td>qPCR (IS1111)</td><td>Vaginal swabs</td><td>Dogs</td><td>0/3</td><td>0%</td><td></td><td>No</td></tr><tr><td>Marié</td><td>NA</td><td>2016</td><td>French</td><td>French Guiana</td><td>Cayenne</td><td>IFI</td><td>Serum</td><td>Dogs</td><td>6–18/158</td><td>3.8%</td><td>6 CT &lt;35</td><td>Yes</td></tr><tr><td>Wei [135]</td><td>2012</td><td>2014</td><td>English</td><td>Nicaragua</td><td>Rivas, South-western Nicaragua</td><td>qPCR</td><td>Whole serum</td><td>Cats</td><td>0/10</td><td>0%</td><td>12 CT &gt;35</td><td>No</td></tr><tr><td>Somma-Moreira [115, 136, 137]</td><td>1956</td><td>1987</td><td>Spanish</td><td>Uruguay</td><td>–</td><td>CAT and CFT</td><td>Serum</td><td>Guinea pigs</td><td>0/31</td><td>0%</td><td>Unpublished data</td><td>No</td></tr></tbody></table>

CAT, capillary agglutination test; CFT, complement fixation test; MA, microagglutination test; ELISA, enzyme-linked immunosorbent assay; IFA, indirect immunofluorescence assay; IFI, indirect immunofluorescence assay; PCR, qualitative polymerase chain reaction; qPCR, quantitative real-time polymerase chain reaction; MLVA, multilocus variable number tandem repeats analysis; MST, multispancer sequence type; IHC, immunohistochemistry; NA, data not available
<table><caption>Table 2 (continued)</caption><thead><tr><th>Author</th><th>Year of the study</th><th>Year of publication</th><th>Language</th><th>Country</th><th>State</th><th>Biological methodology</th><th>Sample type</th><th>Species</th><th>Number of samples</th><th>Result</th><th>Comments</th><th>Published</th></tr></thead><tbody><tr><td>Debin [108]</td><td>2007</td><td>2007</td><td>French</td><td>French Guiana</td><td>Communes du littoral</td><td>ELISA</td><td>Serum</td><td>Dogs</td><td>16/63 (including 8 doubtful cases, 4 positive animals, and 2 positive animals belonging to the same owner</td><td>9.2 to 21.7%</td><td>22 from Kourou kennels and 41 from veterinarians</td><td>Yes</td></tr><tr><td>Davoust</td><td>2013</td><td>2014</td><td>French</td><td>French Guiana</td><td>Montagne du Tigre, Cayenne</td><td>qPCR (IS1111)</td><td>Vaginal swabs</td><td>Dogs</td><td>0/3</td><td>0%</td><td>Unpublished data</td><td>No</td></tr><tr><td>Davoust</td><td>2013</td><td>2014</td><td>French</td><td>French Guiana</td><td>IFA</td><td>IFA</td><td>Serum</td><td>Cat</td><td>0/1</td><td>0%</td><td></td><td></td></tr><tr><td>Davoust [109]</td><td>2013</td><td>2014</td><td>English</td><td>French Guiana</td><td>Cayenne and Kourou</td><td>qPCR (IS1111)</td><td>Vaginal swabs</td><td>Dogs</td><td>0/3</td><td>0%</td><td></td><td>No</td></tr><tr><td>Marié</td><td>NA</td><td>2016</td><td>French</td><td>French Guiana</td><td>Cayenne</td><td>IFI</td><td>Serum</td><td>Dogs</td><td>6–18/158</td><td>3.8%</td><td>6 CT &lt;35</td><td>Yes</td></tr><tr><td>Wei [135]</td><td>2012</td><td>2014</td><td>English</td><td>Nicaragua</td><td>Rivas, South-western Nicaragua</td><td>qPCR</td><td>Whole serum</td><td>Cats</td><td>0/10</td><td>0%</td><td>12 CT &gt;35</td><td>No</td></tr><tr><td>Somma-Moreira [115, 136, 137]</td><td>1956</td><td>1987</td><td>Spanish</td><td>Uruguay</td><td>—</td><td>CAT and CFT</td><td>Serum</td><td>Guinea pigs</td><td>0/31</td><td>0%</td><td>Unpublished data</td><td>No</td></tr></tbody></table>

CAT, capillary agglutination test; CFT, complement fixation test; MA, microagglutination test; ELISA, enzyme-linked immunosorbent assay; IFA, indirect immunofluorescence assay; IFI, indirect immunofluorescence assay; PCR, qualitative polymerase chain reaction; qPCR, quantitative real-time polymerase chain reaction; MLVA, multilocus variable number tandem repeats analysis; MST, multispancer sequence type; IHC, immunohistochemistry; NA, data not available

<figure><img src="image_15.png" /></figure>
<figure><img src="image_16.png" /><figcaption>**Fig. 3** Map of the publications on *Coxiella burnetii* infection in pets in Latin America and the Caribbean</figcaption></figure>

Q fever situation in French Guiana is unique, with the highest incidence in the world, with a peak in the mid-2000s at more than 150 cases per 100,000 inhabitants per year and a stabilization since 2009 of between 25 and 35 cases per 100,000 inhabitants per year, which is much higher than in the rest of Latin America where the disease is barely reported, making French Guiana a hyper-endemic zone [28••, 55, 67]. For comparison, Q fever incidence in mainland France was 1.9/100,000 inhabitants in 2008–2011, and the incidence reached 69/100,000 inhabitants per year in the Netherlands during the largest outbreak ever in 2009, in the population living within 5 km of an infected dairy goat farm, and 6/100,000 inhabitants per year beyond that distance [21, 196]. The distribution of cases is heterogeneous and is mainly concentrated in the capital, Cayenne, and its surroundings, although a recent seroprevalence study suggests that the infection may be more widely distributed across the country than previously reported [197]. A unique strain was identified a few years ago, found exclusively in French Guiana, MST17, both in humans and animals, when the strain was sequenced [59, 109]. One hypothesis is that the incidence of Q fever in humans is “over-reported” in French Guiana compared to other countries because the more virulent MST17 strain causes a less asymptomatic disease (with atypical symptoms) and that Guianese is therefore “over-diagnosed” compared to other countries.

While lung disease usually represents 30 to 50% of cases, it affects more than 90% of patients in this French territory [21, 57]. Finally, one of the most questioning specificities to date of coxiellosis in French Guiana is its animal reservoir, which seems to contrast sharply with the rest of the world. Concerning domestic ruminants, usually considered the main source of transmission of the pathogen to humans, two serological surveys were carried out in the 1990s and 2000, respectively, suggesting a low seroprevalence in ruminants [107, 108, 152]. Ad hoc investigations on farms have been systematically negative, including investigations around cases of Q fever in farmers. The occurrence of several outbreaks of Q fever on the outskirts of Cayenne had raised the possibility of the bacterium being transported by sea winds from certain nearby herds, but this hypothesis has to be corroborated. That is why research has shifted for more than 20 years away from livestock to focus on the search for the bacterium in wildlife surrounding human cases [109, 151, 154]. Nowadays, several individuals from various species of nonflying mammals and bats have been found positive (Table 4), although mostly with Ct > 35, which means that the *C. burnetii* load is detectable but below the limit of quantification. We do not exactly know the link between these species and transmission to humans, apart from the positive sloth incriminated in the epidemic among the military families of Tiger Hill [109, 151]. Moreover, with

<figure><img src="image_17.png" /></figure>
<figure><img src="image_16.png" /><figcaption>**Fig. 3** Map of the publications on *Coxiella burnetii* infection in pets in Latin America and the Caribbean</figcaption></figure>

Q fever situation in French Guiana is unique, with the highest incidence in the world, with a peak in the mid-2000s at more than 150 cases per 100,000 inhabitants per year and a stabilization since 2009 of between 25 and 35 cases per 100,000 inhabitants per year, which is much higher than in the rest of Latin America where the disease is barely reported, making French Guiana a hyper-endemic zone [28••, 55, 67]. For comparison, Q fever incidence in mainland France was 1.9/100,000 inhabitants in 2008–2011, and the incidence reached 69/100,000 inhabitants per year in the Netherlands during the largest outbreak ever in 2009, in the population living within 5 km of an infected dairy goat farm, and 6/100,000 inhabitants per year beyond that distance [21, 196]. The distribution of cases is heterogeneous and is mainly concentrated in the capital, Cayenne, and its surroundings, although a recent seroprevalence study suggests that the infection may be more widely distributed across the country than previously reported [197]. A unique strain was identified a few years ago, found exclusively in French Guiana, MST17, both in humans and animals, when the strain was sequenced [59, 109]. One hypothesis is that the incidence of Q fever in humans is “over-reported” in French Guiana compared to other countries because the more virulent MST17 strain causes a less asymptomatic disease (with atypical symptoms) and that Guianese is therefore “over-diagnosed” compared to other countries.

While lung disease usually represents 30 to 50% of cases, it affects more than 90% of patients in this French territory [21, 57]. Finally, one of the most questioning specificities to date of coxiellosis in French Guiana is its animal reservoir, which seems to contrast sharply with the rest of the world. Concerning domestic ruminants, usually considered the main source of transmission of the pathogen to humans, two serological surveys were carried out in the 1990s and 2000, respectively, suggesting a low seroprevalence in ruminants [107, 108, 152]. Ad hoc investigations on farms have been systematically negative, including investigations around cases of Q fever in farmers. The occurrence of several outbreaks of Q fever on the outskirts of Cayenne had raised the possibility of the bacterium being transported by sea winds from certain nearby herds, but this hypothesis has to be corroborated. That is why research has shifted for more than 20 years away from livestock to focus on the search for the bacterium in wildlife surrounding human cases [109, 151, 154]. Nowadays, several individuals from various species of nonflying mammals and bats have been found positive (Table 4), although mostly with Ct > 35, which means that the *C. burnetii* load is detectable but below the limit of quantification. We do not exactly know the link between these species and transmission to humans, apart from the positive sloth incriminated in the epidemic among the military families of Tiger Hill [109, 151]. Moreover, with

<figure><img src="image_17.png" /></figure>
<figure><img src="image_18.png" /><figcaption>**Fig. 4** Map of the publications on *Coxiella burnetii* infection in wildlife in Latin America and the Caribbean</figcaption></figure>

the exception of the military clusters reported earlier, the numerous cases of Q fever reported in French Guiana each year are generally isolated cases, without any clustering of cases, which does not facilitate the understanding of the origin of the transmission. One possibility would be the persistence of the bacterium in the environment as a result of different wildlife species emitting it, and that humans could become infected by inhalation of this dust as a result of favorable circumstances like leveling work near the house, gardening, particularly the use of a brush cutter, sweeping, etc. [57, 107, 154].

Recently, two studies have challenged the dogma that there is no link between livestock and Q fever cases in Guiana. On one hand, Saout et al. showed that cattle had a high seroprevalence of Q fever, especially when based not on individual prevalence but on herd prevalence [122]. On the other hand, seroprevalence study for *C. burnetii* of almost 3000 people sampled throughout the country showed that there was a link between high prevalence and proximity to livestock farms [197]. To explain these contradictory results, the old studies focused on a small number of animals and/or herds. In addition, studies conducted in the 1990s used the complement fixation test, currently considered to be weakly sensitive [198]. Moreover, several cases of Q fever were reported among workers at the Cayenne abattoir in 2007

[108]. New studies are therefore needed in French Guiana on livestock to more strictly support these new hypotheses.

In conclusion, the epidemiological cycle of Q fever in French Guiana is still unresolved. Although several species of wild mammals have been identified as carriers of *C. burnetii*, these results remain anecdotal and explain neither the frequency of the disease nor the geographical and ethnic distribution of cases in the territory. Moreover, the dogma that *C. burnetii* was not transmitted by cattle in French Guiana seems questionable. Additional investigations of the animal reservoir, livestock, and wildlife must be carried out to allow a better understanding of the unique phenomenon observed in this small piece of Amazonia.

## Brazil

In Brazil, where the first reports of *C. burnetii* infection occurred in the 1950s through serological tests in human and cattle samples [199], Q fever is currently subject to compulsory notification in the context of the differential diagnosis of rickettsioses. However, official data are lacking, which can impact the understanding of the epidemiology of this zoonosis in the country. Furthermore, although some human cases of Q fever have been mandatorily reported, the high incidence of febrile illnesses such as malaria, leptospirosis, dengue, Zika, and chikungunya, can lead to misdiagnosis of Q fever [66].
<figure><img src="image_18.png" /><figcaption>**Fig. 4** Map of the publications on *Coxiella burnetii* infection in wildlife in Latin America and the Caribbean</figcaption></figure>

the exception of the military clusters reported earlier, the numerous cases of Q fever reported in French Guiana each year are generally isolated cases, without any clustering of cases, which does not facilitate the understanding of the origin of the transmission. One possibility would be the persistence of the bacterium in the environment as a result of different wildlife species emitting it, and that humans could become infected by inhalation of this dust as a result of favorable circumstances like leveling work near the house, gardening, particularly the use of a brush cutter, sweeping, etc. [57, 107, 154].

Recently, two studies have challenged the dogma that there is no link between livestock and Q fever cases in Guiana. On one hand, Saout et al. showed that cattle had a high seroprevalence of Q fever, especially when based not on individual prevalence but on herd prevalence [122]. On the other hand, seroprevalence study for *C. burnetii* of almost 3000 people sampled throughout the country showed that there was a link between high prevalence and proximity to livestock farms [197]. To explain these contradictory results, the old studies focused on a small number of animals and/or herds. In addition, studies conducted in the 1990s used the complement fixation test, currently considered to be weakly sensitive [198]. Moreover, several cases of Q fever were reported among workers at the Cayenne abattoir in 2007

[108]. New studies are therefore needed in French Guiana on livestock to more strictly support these new hypotheses.

In conclusion, the epidemiological cycle of Q fever in French Guiana is still unresolved. Although several species of wild mammals have been identified as carriers of *C. burnetii*, these results remain anecdotal and explain neither the frequency of the disease nor the geographical and ethnic distribution of cases in the territory. Moreover, the dogma that *C. burnetii* was not transmitted by cattle in French Guiana seems questionable. Additional investigations of the animal reservoir, livestock, and wildlife must be carried out to allow a better understanding of the unique phenomenon observed in this small piece of Amazonia.

## Brazil

In Brazil, where the first reports of *C. burnetii* infection occurred in the 1950s through serological tests in human and cattle samples [199], Q fever is currently subject to compulsory notification in the context of the differential diagnosis of rickettsioses. However, official data are lacking, which can impact the understanding of the epidemiology of this zoonosis in the country. Furthermore, although some human cases of Q fever have been mandatorily reported, the high incidence of febrile illnesses such as malaria, leptospirosis, dengue, Zika, and chikungunya, can lead to misdiagnosis of Q fever [66].
<figure><img src="image_19.png" /></figure>

<table><caption>**Table 3** Studies on *Coxiella burnetii* in wildlife in Latin America</caption><thead><tr><th>Author</th><th>Year of the study</th><th>Year of publication</th><th>Language</th><th>Country</th><th>Location</th><th>Methodology</th><th>Sample type</th><th>Species</th><th>Number of samples</th><th>Result</th><th>Comments</th><th>Published</th></tr></thead><tbody><tr><td>Mares-Guia [77]</td><td>2011–2012</td><td>2016</td><td>Portuguese</td><td>Brazil</td><td>Município de Itaborai, Rio de Janeiro State</td><td>PCR (htpAB)</td><td>Spleen</td><td>Rodents and marsupials*</td><td>0/60</td><td>0%</td><td>Unpublished thesis</td><td>No</td></tr><tr><td>Rozental [139]</td><td>2007–2012</td><td>2017</td><td>English</td><td>Brazil</td><td>8 municipalities of Rio de Janeiro State (Atlantic forest)</td><td>PCR (IS1111)</td><td>Spleen</td><td>Rodents<br/>Akodon cursor [3]<br/>M. musculus [1]<br/>O. dasytrichus [1]<br/>O. nigripes [1]</td><td>6/131</td><td>4.6%</td><td></td><td>Yes</td></tr><tr><td>Ferreira [140]</td><td>2013–2015</td><td>2018</td><td>English</td><td>Brazil</td><td>Bahia, Rio de Janeiro and Santa Catarina States (Atlantic forest)</td><td>PCR (IS1111)</td><td>Spleen</td><td>Bats<br/>A. lituratus [3]<br/>A. fimbriatus [1]</td><td>4/119</td><td>3.4%</td><td></td><td>Yes</td></tr><tr><td>Zanatto [141]</td><td>1996–2011</td><td>2019</td><td>English</td><td>Brazil</td><td>Mato Grasso do Sul, São Paulo, Goiás and Paraná</td><td>PCR (IS1111)</td><td>Serum</td><td>Free-living civids*</td><td>0/188</td><td>0%</td><td></td><td>Yes</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td>IFA</td><td></td><td>Blastocerus dichotomus<br/>M. gouazoubira</td><td>9/169</td><td>5.3%</td><td></td><td></td></tr><tr><td>Colle [142]</td><td>2014</td><td>2019</td><td>English</td><td>Brazil</td><td>Alta Floresta, Sinop and Cláudia, Mato Grosso (Amazonian biome)</td><td>PCR Coxiella CTP synthase (pyrG)</td><td>Liver, spleen, serum</td><td>Rodents $<br/>Marsupials $</td><td>0/78<br/>0/152</td><td>0%<br/>0%</td><td></td><td>Yes</td></tr><tr><td>Barboza de Oliveira [131]</td><td>2014–2016</td><td>2020</td><td>English</td><td>Brazil</td><td>Ceará and Pernambuco (Caatinga biome)</td><td>IFA</td><td>Serum</td><td>Rodents (Wiedomys pyrrohylinos)<br/>Marsupials (Didelphis albiventris)<br/>Wild canids*</td><td>1/51<br/>1/18<br/>0/1<br/>0/51<br/>0/18<br/>0/1<br/>0/60</td><td>2%<br/>5.6%<br/>0%<br/>0%<br/>0%<br/>0%<br/>0%</td><td></td><td>Yes</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td>PCR</td><td></td><td>Marsupials<br/>Wild canids</td><td>121 reptiles<br/>49 amphibians<br/>60 samples tested for pathogens</td><td>0/60<br/>0%</td><td></td><td>Yes</td></tr><tr><td>Mendoz-Roldan [143]</td><td>2015–2018</td><td>2020</td><td>English</td><td>Brazil</td><td>States: Acre, Espirito Santo, Mato Grosso, Mato Grosso do Sul, Minas Gerais, Pará, Rio de Janeiro and São Paulo</td><td>Conventional PCR (Coxiella spp.—capsular polysaccharide biosynthesis protein I-like gene (cap))</td><td>48 serum<br/>12 livers</td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>Ikeda [144]</td><td>2017–201</td><td>2021</td><td>English</td><td>Brazil</td><td>Campo Grande city, Mato Grosso do Sul state</td><td>qPCR (IS1111)</td><td>133 serum<br/>135 spleen</td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>Santana [145]</td><td>2018–2020</td><td>2022</td><td>English</td><td>Brazil</td><td>Municipalities of Barretos, Colina, Guaraci, Jabotibaçal, Monte Alto, Monte Azul<br/>Paulista, Morro Agudo, Olímpia, and Torrinha, São Paulo State</td><td>qPCR (IS1111)</td><td>Serum</td><td>Free-living wild boars (Sus scrofa)</td><td>135 nonheumatophagous bats*</td><td>0/135</td><td>0%</td><td></td><td>Yes</td></tr></tbody></table>
<table><caption>**Table 3** Studies on *Coxiella burnetii* in wildlife in Latin America</caption><thead><tr><th>Author</th><th>Year of the study</th><th>Year of publication</th><th>Language</th><th>Country</th><th>Location</th><th>Methodology</th><th>Sample type</th><th>Species</th><th>Number of samples</th><th>Result</th><th>Comments</th><th>Published</th></tr></thead><tbody><tr><td>Mares-Guia [77]</td><td>2011–2012</td><td>2016</td><td>Portuguese</td><td>Brazil</td><td>Município de Itaborai, Rio de Janeiro State</td><td>PCR (htpAB)</td><td>Spleen</td><td>Rodents and marsupials*</td><td>0/60</td><td>0%</td><td>Unpublished thesis</td><td>No</td></tr><tr><td>Rozental [139]</td><td>2007–2012</td><td>2017</td><td>English</td><td>Brazil</td><td>8 municipalities of Rio de Janeiro State (Atlantic forest)</td><td>PCR (IS1111)</td><td>Spleen</td><td>Rodents</td><td>6/131</td><td>4.6%</td><td></td><td>Yes</td></tr><tr><td>Ferreira [140]</td><td>2013–2015</td><td>2018</td><td>English</td><td>Brazil</td><td>Bahia, Rio de Janeiro and Santa Catarina States (Atlantic forest)</td><td>PCR (IS1111)</td><td>Spleen</td><td>Bats</td><td>4/119</td><td>3.4%</td><td></td><td>Yes</td></tr><tr><td>Zanatto [141]</td><td>1996–2011</td><td>2019</td><td>English</td><td>Brazil</td><td>Mato Grasso do Sul, São Paulo, Goiás and Paraná</td><td>PCR (IS1111)</td><td>Serum</td><td>Free-living civids*</td><td>0/188</td><td>0%</td><td></td><td>Yes</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td>IFA</td><td></td><td>Blastocerus dichotomus</td><td>9/169</td><td>5.3%</td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>M. gouazoubira</td><td>7/129</td><td>5.4%</td><td></td><td></td></tr><tr><td>Colle [142]</td><td>2014</td><td>2019</td><td>English</td><td>Brazil</td><td>Alta Floresta, Sinop and Cláudia, Mato Grosso (Amazonian biome)</td><td>PCR Coxiella CTP synthase (pyrG)</td><td>Liver, spleen, serum</td><td>Rodents $</td><td>0/78</td><td>0%</td><td></td><td>Yes</td></tr><tr><td>Barboza de Oliveira [131]</td><td>2014–2016</td><td>2020</td><td>English</td><td>Brazil</td><td>Ceará and Pernambuco (Caatinga biome)</td><td>IFA</td><td>Serum</td><td>Marsupials $</td><td>0/152</td><td>0%</td><td></td><td>Yes</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Rodents (Wiedomys pyrrohylinos)</td><td>1/51</td><td>2%</td><td></td><td>Yes</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Marsupials (Didelphis albiventris)</td><td>1/18</td><td>5.6%</td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Wild canids*</td><td>0/1</td><td>0%</td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Rodents</td><td>0/51</td><td>0%</td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Marsupials</td><td>0/18</td><td>0%</td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Wild canids</td><td>0/1</td><td>0%</td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>121 reptiles</td><td>0/60</td><td>0%</td><td></td><td>Yes</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>49 amphibians</td><td></td><td></td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>60 samples tested for pathogens</td><td></td><td></td><td></td><td></td></tr><tr><td>Mendozaro Roldan [143]</td><td>2015–2018</td><td>2020</td><td>English</td><td>Brazil</td><td>States: Acre, Espirito Santo, Mato Grosso, Mato Grosso do Sul, Minas Gerais, Pará, Rio de Janeiro and São Paulo</td><td>PCR (Coxiella spp.—capsular polysaccharide biosynthesis protein I-like gene (cap))</td><td>Conventional PCR (Coxiella spp.—capsular polysaccharide biosynthesis protein I-like gene (cap))</td><td>48 serum</td><td></td><td></td><td></td><td></td></tr><tr><td>Ikeda [144]</td><td>2017–201</td><td>2021</td><td>English</td><td>Brazil</td><td>Campo Grande city, Mato Grosso do Sul state</td><td>qPCR (IS1111)</td><td>133 serum</td><td>12 livers</td><td></td><td></td><td></td><td></td></tr><tr><td>Santana [145]</td><td>2018–2020</td><td>2022</td><td>English</td><td>Brazil</td><td>Municipalities of Barretos, Colina, Guaraci, Jabotiba cabal, Monte Alto, Monte Azul, Paulista, Morro Agudo, Olímpia, and Torrinha, São Paulo State</td><td>qPCR (IS1111)</td><td>Serum</td><td>135 spleen</td><td></td><td></td><td></td><td></td></tr></tbody></table>

<figure><img src="image_19.png" /></figure>
<table><caption>**Table 3 (continued)**</caption><thead><tr><th>Author</th><th>Year of the study</th><th>Year of publication</th><th>Language</th><th>Country</th><th>Location</th><th>Methodology</th><th>Sample type</th><th>Species</th><th>Number of samples</th><th>Result</th><th>Comments</th><th>Published</th></tr></thead><tbody><tr><td>De Oliveira [146]</td><td>2015–2020</td><td>2022</td><td>English</td><td>Brazil</td><td>Mato Grosso do Sul, São Paulo, Pará, Rondônia and Rio Grande do Sul</td><td>qPCR (ISI/111)</td><td>Serum</td><td>Sloths*</td><td>0/233</td><td>0%</td><td>Samples obtained during the necropsy of animals</td><td>Yes</td></tr><tr><td>Muller [147•]</td><td>2017–2018</td><td>2020</td><td>English</td><td>Chile</td><td>Metropolitan region The Biobio Region La Araucanía Regions</td><td>qPCR (ISI/111)</td><td>Spleen for T. brasiliensis Serum for the others</td><td>Bats<br>Tadarida brasiliensis</td><td>5/55<br>5/9</td><td>9%<br>56%</td><td>Victims of a vehicular collision on highways<br>Quantitation cycle &lt;37<br>All positives from the metropolitan region</td><td>Yes</td></tr><tr><td>Di Cataldo [133]</td><td>2006–2019</td><td>2022</td><td>English</td><td>Chile</td><td>4 different areas: Coastal Desert, Mountain Desert, Steppe-Mediterranean, and Temperate Warm Rainy</td><td>IFLA<br>ELISA</td><td>Blood</td><td>12 South American grey foxes (Lycalopex griseus), 10 Andean foxes (L. culpaeus) and 10 Darwin's foxes</td><td>0/32</td><td>0%</td><td>Yes</td><td>Yes</td></tr><tr><td>Hidalgo-Hermoso [148]</td><td>2013–2018</td><td>2022</td><td>English</td><td>Chile</td><td>Southern Chile</td><td>Serology NA</td><td>Serum</td><td>Darwin's fox (Lycalopex fulvipes)</td><td>0/47</td><td>0%</td><td>Yes</td><td>Yes</td></tr><tr><td>Cabello [149]</td><td>2009–2012</td><td>2013</td><td>English</td><td>Chile</td><td>Chiloé Island</td><td>qPCR</td><td>Serum</td><td>Darwin's fox (Lycalopex fulvipes)</td><td>0/30</td><td>0%</td><td>Yes</td><td>Yes</td></tr><tr><td>Silva-Ramos [150•]</td><td>201–2015, 2018</td><td>2022</td><td>English</td><td>Colombia</td><td>Macaregua cave, Las Vueltas village, Municipality of Curitii, Santander Department</td><td>qPCR ISI111</td><td>Serum</td><td>Bats<br>C. perspicillata<br>M. megatophylla<br>N. tumidirostris</td><td>8/126<br>3/49<br>2/35<br>3/42</td><td>6.3%<br>6.1%<br>5.7%<br>7.1%</td><td>Yes</td><td>Yes</td></tr></tbody></table>
<table><caption>Table 3 (continued)</caption><thead><tr><th>Author</th><th>Year of the study</th><th>Year of publication</th><th>Language</th><th>Country</th><th>Location</th><th>Methodology</th><th>Sample type</th><th>Species</th><th>Number of samples</th><th>Result</th><th>Comments</th><th>Published</th></tr></thead><tbody><tr><td>De Oliveira [146]</td><td>2015–2020</td><td>2022</td><td>English</td><td>Brazil</td><td>Mato Grosso do Sul, São Paulo, Pará, Rondônia and Rio Grande do Sul</td><td>qPCR (ISI/111)</td><td>Serum</td><td>Sloths*</td><td>0/233</td><td>0%</td><td>Samples obtained during the necropsy of animals</td><td>Yes</td></tr><tr><td>Muller [147•]</td><td>2017–2018</td><td>2020</td><td>English</td><td>Chile</td><td>Metropolitan region The Biobio Region La Araucanía Regions</td><td>qPCR (ISI/111)</td><td>Spleen for T. brasiliensis Serum for the others</td><td>Bats Tadarida brasiliensis</td><td>5/55<br>5/9</td><td>9%<br>56%</td><td>Victims of a vehicular collision on highways<br>Quantitation cycle &lt;37<br>All positives from the metropolitan region</td><td>Yes</td></tr><tr><td>Di Cataldo [133]</td><td>2006–2019</td><td>2022</td><td>English</td><td>Chile</td><td>4 different areas: Coastal Desert, Mountain Desert, Steppe-Mediterranean, and Temperate Warm Rainy</td><td>IFLA ELISA</td><td>Blood</td><td>12 South American grey foxes (Lycalopex griseus), 10 Andean foxes (L. culpaeus) and 10 Darwin's foxes</td><td>0/32</td><td>0%</td><td>Yes</td><td>Yes</td></tr><tr><td>Hidalgo-Hermoso [148]</td><td>2013–2018</td><td>2022</td><td>English</td><td>Chile</td><td>Southern Chile</td><td>Serology NA</td><td>Serum</td><td>Darwin's fox (Lycalopex fulvipes)</td><td>0/47</td><td>0%</td><td>Yes</td><td>Yes</td></tr><tr><td>Cabello [149]</td><td>2009–2012</td><td>2013</td><td>English</td><td>Chile</td><td>Chiloé Island</td><td>qPCR</td><td>Serum</td><td>Darwin's fox (Lycalopex fulvipes)</td><td>0/30</td><td>0%</td><td>Yes</td><td>Yes</td></tr><tr><td>Silva-Ramos [150•]</td><td>201–2015, 2018</td><td>2022</td><td>English</td><td>Colombia</td><td>Macaregua cave, Las Vueltas village, Municipality of Curitii, Santander Department</td><td>qPCR ISI111</td><td>Serum</td><td>Bats C. perspicillata M. megatophylla N. tumidirostris</td><td>8/126<br>3/49<br>2/35<br>3/42</td><td>6.3%<br>6.1%<br>5.7%<br>7.1%</td><td>Yes</td><td>Yes</td></tr></tbody></table>

<figure><img src="image_20.png" /></figure>
<table><caption>**Table 3 (continued)**</caption><thead><tr><th>Author</th><th>Year of the study</th><th>Year of publication</th><th>Language</th><th>Country</th><th>Location</th><th>Methodology</th><th>Sample type</th><th>Species</th><th>Number of samples</th><th>Result</th><th>Comments</th><th>Published</th></tr></thead><tbody><tr><td>Gardon [107]</td><td>1998–2000</td><td>2001</td><td>English</td><td>French Guiana</td><td>Cayenne and surroundings</td><td>IFA</td><td>Serum</td><td>Rodents<br>Proechimys sp.</td><td>4/117<br>4/26</td><td>3.4%<br>15.4%</td><td>Captured in various areas near the houses of case patients with Q fever</td><td>Yes</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Marsupials<br>P. opossum</td><td>5/42<br>4/36</td><td>11.9%<br>11.1%</td><td>Swallows were captured on the roof of the jail at Remire, where many cases of Q fever had been described among guards and prisoners</td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Bats<br>D.marsupialis</td><td>1/4<br>0/86</td><td>25%<br>0%</td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Birds<br>Batrachians</td><td>1/69<br>0/47</td><td>0%<br>1.4%</td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Rodents<br>Marsupials</td><td>0/42<br>0/17</td><td>0%<br>0%</td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Bats<br>Batrachians</td><td>0/42<br>0/69</td><td>0%<br>0%</td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Lung, liver, or intestine</td><td>0/42<br>0/69</td><td>0%<br>0%</td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>PCR</td><td>0/86<br>0/42</td><td>0%<br>0%</td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>Davoust [109]</td><td>2013–2014</td><td>2014</td><td>English</td><td>French Guiana</td><td>Montagne du tigre, Cayenne</td><td>qPCR (IS1111)</td><td>Feces, spleen</td><td>3-toed sloth (female) (Brady-pus tridactylus)</td><td>1/1<br>0/7</td><td>100%<br>0%</td><td></td><td>Yes</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Vaginal swabs</td><td>0/34<br>0/2</td><td>0%<br>0%</td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Opossum<br>Iguana</td><td>0/4<br>0/17</td><td>0%<br>0%</td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Gecko (species not available)</td><td>0/17<br>0/12</td><td>0%<br>16.6%</td><td></td><td></td></tr><tr><td>Pommier de Santi [151]</td><td>2013</td><td>2014</td><td>English</td><td>French Guiana</td><td>Montagne du tigre, Cayenne</td><td>IFA</td><td>Serum</td><td>Didelphis marsupialis<br>Philander opossum</td><td>2/12<br>1/6</td><td>16.6%<br>16.6%</td><td></td><td>Yes</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Didelphis marsupialis</td><td>0/12<br>0/16</td><td>0%<br>0%</td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Serum [14]<br>Philander opossum</td><td>0/6<br>0/16</td><td>0%<br>0%</td><td></td><td></td></tr><tr><td>Pommier de Santi [152]</td><td>2014</td><td>2014</td><td>English</td><td>French Guiana</td><td>Montagne du tigre (Cayenne) and Macouria Sloths Rescue Center</td><td>qPCR (IS1111)</td><td>Vaginal swabs [18]<br>Vaginal and rectal swabs + hairs collected in the perianal area + feces</td><td>Bradypus tridactylus</td><td>0/16<br>0/12</td><td>0%<br>0%</td><td>3 collected on the Montagne du Tigre and 13 were sampled in Macouria</td><td>Yes</td></tr></tbody></table>

<figure><img src="image_20.png" /></figure>
<table><caption>**Table 3 (continued)**</caption><thead><tr><th>Author</th><th>Year of the study</th><th>Year of publication</th><th>Language</th><th>Country</th><th>Location</th><th>Methodology</th><th>Sample type</th><th>Species</th><th>Number of samples</th><th>Result</th><th>Comments</th><th>Published</th></tr></thead><tbody><tr><td>Gardon [107]</td><td>1998–2000</td><td>2001</td><td>English</td><td>French Guiana</td><td>Cayenne and surroundings</td><td>IFA</td><td>Serum</td><td>Rodents<br>Proechimys sp.</td><td>4/117<br>4/26</td><td>3.4%<br>15.4%</td><td>Captured in various areas near the houses of case patients with Q fever</td><td>Yes</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Marsupials<br>P. opossum</td><td>5/42<br>4/36</td><td>11.9%<br>11.1%</td><td>Swallows were captured on the roof of the jail at Remire, where many cases of Q fever had been described among guards and prisoners</td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Bats<br>D.marsupialis</td><td>1/4<br>0/86</td><td>25%<br>0%</td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Birds<br>Batrachians</td><td>1/69<br>0/47</td><td>0%<br>1.4%</td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Rodents<br>Marsupials</td><td>0/42<br>0/17</td><td>0%<br>0%</td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Bats<br>Batrachians</td><td>0/42<br>0/69</td><td>0%<br>0%</td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Lung, liver, or intestine</td><td>0/42<br>0/69</td><td>0%<br>0%</td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>Davoust [109]</td><td>2013–2014</td><td>2014</td><td>English</td><td>French Guiana</td><td>Montagne du tigre, Cayenne</td><td>qPCR (IS1111)</td><td>Feces, spleen</td><td>3-toed sloth (female) (Brady-pus tridactylus)</td><td>1/1</td><td>100%<br>0%</td><td></td><td>Yes</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Vaginal swabs</td><td>0/7<br>0/34</td><td>0%<br>0%</td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Bats<br>Opossum<br>Iguana</td><td>0/2<br>0/4</td><td>0%<br>0%</td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Gecko (species not available)</td><td>0/17</td><td>0%</td><td></td><td></td></tr><tr><td>Pommier de Santi [151]</td><td>2013</td><td>2014</td><td>English</td><td>French Guiana</td><td>Montagne du tigre, Cayenne</td><td>IFA</td><td>Serum</td><td>Didelphis marsupialis<br>Philander opossum</td><td>2/12<br>1/6</td><td>16.6%<br>16.6%</td><td></td><td>Yes</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Didelphis marsupialis</td><td>0/12</td><td>0%</td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Serum [14]<br>Philander opossum</td><td>0/6<br>0/16</td><td>0%<br>0%</td><td></td><td></td></tr><tr><td>Pommier de Santi [152]</td><td>2014</td><td>2014</td><td>English</td><td>French Guiana</td><td>Montagne du tigre (Cayenne) and Macouria Sloths Rescue Center</td><td>qPCR (IS1111)</td><td>Vaginal swabs [18]<br>Vaginal and rectal swabs + hairs collected in the perianal area + feces</td><td>Bradypus tridactylus</td><td>0/16</td><td>0%</td><td></td><td>3 collected on the Montagne du Tigre and 13 were sampled in Macouria</td></tr></tbody></table>

<figure><img src="image_21.png" /></figure>
<table><caption>**Table 3 (continued)**</caption><thead><tr><th>Author</th><th>Year of the study</th><th>Year of publication</th><th>Language</th><th>Country</th><th>Location</th><th>Methodology</th><th>Sample type</th><th>Species</th><th>Number of samples</th><th>Result</th><th>Comments</th><th>Published</th></tr></thead><tbody><tr><td>Marié [153]</td><td>2005–2006</td><td>2013</td><td>French</td><td>French Guiana</td><td></td><td>qPCR (IS111)</td><td>Muscular juice</td><td>Myoproeta acouchi Tayassu pecari</td><td>0/1<br>3/35</td><td>0%<br>8.6%</td><td>3 with CT > 35<br>1 with CT > 35/l<br>with CT < 35</td><td>No</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Hydrochoerus hydrochaeris</td><td>2/30</td><td>6.7%</td><td>1 with CT > 35/l<br>with CT < 35</td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Tapirus terrestris</td><td>2/7</td><td>28.6%</td><td>3 with CT > 35</td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Pecari tajacu</td><td>3/13</td><td>23.0%</td><td>1 with CT > 35</td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Dasypus novemcinctus</td><td>1/5</td><td>20.0%</td><td>1 with CT > 35</td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Liver</td><td>Rodents*</td><td>1/30</td><td>3%</td><td>Unpublished data</td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Prochimys cavieri</td><td>1/4</td><td>25%</td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Bats*</td><td>6/199</td><td>3.0%</td><td>6 pos but with CT > 35</td><td>No</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>N. albiventris</td><td>1/14</td><td></td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>P. parnellii</td><td>1/37</td><td></td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>C. perspicillata</td><td>3/76</td><td>0%</td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>P. elongatus</td><td>1/13</td><td></td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Serum</td><td>Bats</td><td>0/41</td><td>0%</td><td>No</td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Species not available</td><td></td><td></td><td></td><td></td></tr><tr><td>Davoust [153]</td><td>2013</td><td>2013</td><td>French</td><td>French Guiana</td><td>Cayenne, Saint Laurent du Maroni, Kourou</td><td>qPCR (IS30.A)</td><td>Vaginal swab samples</td><td>Capybara (Hydrochoerus hydrochaeris)</td><td>1/1</td><td>100%</td><td>MST 17</td><td>Yes</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Capybara (Hydrochoerus hydrochaeris)</td><td></td><td></td><td></td><td></td></tr><tr><td>Davoust</td><td>2013 and 2014</td><td></td><td>French</td><td>French Guiana</td><td>Cayenne, Saint Laurent du Maroni, Kourou</td><td>ELISA</td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>Christen [154]</td><td>2014</td><td>2019</td><td>English</td><td>French Guiana</td><td>La Comté river, Roura</td><td>qPCR (IS111)</td><td>Stool</td><td></td><td></td><td></td><td></td><td></td></tr></tbody></table>

<figure><img src="image_21.png" /></figure>
<table><caption>Table 3 (continued)</caption><thead><tr><th>Author</th><th>Year of the study</th><th>Year of publication</th><th>Language</th><th>Country</th><th>Location</th><th>Methodology</th><th>Sample type</th><th>Species</th><th>Number of samples</th><th>Result</th><th>Comments</th><th>Published</th></tr></thead><tbody><tr><td>Marié [153]</td><td>2005–2006</td><td>2013</td><td>French</td><td>French Guiana</td><td></td><td>qPCR (IS111)</td><td>Muscular juice</td><td>Myoproeta acouchi Tayassu pecari</td><td>0/1<br>3/35</td><td>0%<br>8.6%</td><td>3 with CT > 35<br>1 with CT > 35/l<br>with CT < 35</td><td>No</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Hydrochoerus hydrochaeris</td><td>2/30</td><td>6.7%</td><td>1 with CT > 35/l<br>with CT < 35</td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Tapirus terrestris</td><td>2/7</td><td>28.6%</td><td>3 with CT > 35</td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Pecari tajacu</td><td>3/13</td><td>23.0%</td><td>1 with CT > 35</td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Dasypus novemcinctus</td><td>1/5</td><td>20.0%</td><td>1 with CT > 35</td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Liver</td><td>Rodents*</td><td>1/30</td><td>3%</td><td>Unpublished data</td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Proechimys caviieri</td><td>1/4</td><td>25%</td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Bats*</td><td>6/199</td><td>3.0%</td><td>6 pos but with CT > 35</td><td>No</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>N. albiventris</td><td>1/14</td><td></td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>P. parnellii</td><td>1/37</td><td></td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>C. perspicillata</td><td>3/76</td><td>0%</td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>P. elongatus</td><td>1/13</td><td></td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Serum</td><td>Bats</td><td>0/41</td><td>0%</td><td>No</td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Species not available</td><td></td><td></td><td></td><td></td></tr><tr><td>Davoust [153]</td><td>2013</td><td>2013</td><td>French</td><td>French Guiana</td><td>Cayenne, Saint Laurent du Maroni, Kourou</td><td>qPCR (IS30.A)</td><td>Vaginal swab samples</td><td>Capybara (Hydrochoerus hydrochaeris)</td><td>1/1</td><td>100%</td><td>MST 17</td><td>Yes</td></tr><tr><td>Davoust</td><td>2013 and 2014</td><td>2019</td><td>English</td><td>French Guiana</td><td>Cayenne, Saint Laurent du Maroni, Kourou</td><td>ELISA</td><td></td><td>Stool</td><td></td><td></td><td></td><td></td></tr><tr><td>Christen [154]</td><td>2014</td><td>2019</td><td>English</td><td>French Guiana</td><td>La Comté river, Roura</td><td>qPCR (IS111)</td><td></td><td></td><td></td><td></td><td></td><td></td></tr></tbody></table>
<table><caption>**Table 3** (continued)</caption><thead><tr><th>Author</th><th>Year of the study</th><th>Year of publication</th><th>Language</th><th>Country</th><th>Location</th><th>Methodology</th><th>Sample type</th><th>Species</th><th>Number of samples</th><th>Result</th><th>Comments</th><th>Published</th></tr></thead><tbody><tr><td>Destoop</td><td>2018</td><td></td><td>French</td><td>French Guiana</td><td>Cayenne</td><td>qPCR (IS111)</td><td>Stool</td><td>Captive song-birds + +</td><td>0/11</td><td>0%</td><td>Unpublished data</td><td>No</td></tr><tr><td>Somma-Moreira [115, 116]</td><td>1985</td><td>1987</td><td>English and Spanish</td><td>Uruguay</td><td>NA</td><td>MAT</td><td>Serum</td><td>Birds</td><td>0/66</td><td>0%</td><td></td><td>Yes</td></tr><tr><td>Hernandez [155]</td><td>2003–2004</td><td>2007</td><td>English</td><td>Uruguay</td><td>Dpt of Maldonado</td><td>IFA</td><td>Serum</td><td>Pampas deer (Ozotoceros bezoarticus)</td><td>5/22</td><td>22%</td><td>Experimental wildlife breeding station department</td><td>Yes</td></tr></tbody></table>

CFT, complement fixation test; MA, microagglutination test; ELISA, enzyme-linked immunosorbent assay; IFA, immunofluorescence assay; PCR, qualitative polymerase chain reaction; qPCR, quantitative real-time polymerase chain reaction; MLVA, multilocus variable number tandem repeats analysis; MST, multispacer sequence type.; IHC, immunohistochemistry

*Species from Mares-Guia et al. [77]: Rodentia: Akodon cursor (n=10), Nectomys squamipes [11], Oligoryzomys nigripes [11], Didelphimorpha: Didelphis aurita (n=9); Micoureus (marmosa) paraguayanus (n=12); and Philander frenatus (n=27). Rodents species from Rozenthal et al. [139]: Akodon cursor [32], Mus musculus [19], Oligoryzomys nigripes [16], Delomys doralis [14], Oxymycteris dasytrichus [12], Euryoryzomys russipes [7], Trinomys iheringi [6], Nectomys squamipes [5], Akodon lindberghi [5], Trinomys dimidiatus [4], Akodon montensis [3], Oligoryzomys flavescens [2], Brucepatersonius iheringi [1], Calomys tener [1], Necromys lasiurus [1], Oecomys catherinae [1], Oxymycterus delator [1], and Rhipidomys itoan [1]. Bats species from Ferreira et al. [140]: Carollia perspicillata (n=34), Desmodus rotundus [15], Arthebus lituratus [14], Sturnira lilium [12], Arthebus fimbriatus [7], Rhinophylla pumilio [7] Arthebus planirostris [5], Dermanura cinerea [4], Phyllostomus discolor [4], Arthebus obscurus [2], Glossophaga soricina [2], Myotis nigricans [2], Sturnira tildae [2], Vampyressa pusilla [2], Anoura caudifer [1], Chiroderma doriae [1], Lonchophylla peracchii [1], Micronycteris minuta [1], Micronycteris sp. [1], Phyllostomus hastatus [1], and Trinomycteris nicefori [1]. Free-living deer species from Zanatto et al. [141]: Blastocerus dichotomus [156], Mazama gouazoubira [27], M. bororo [4], M. americana [3], and Ozotocerus bezoarticus [11]. Rodents and marsupials species from Colle [142]: rodents: Euryoryzomys nitidus [1], Hylaeamys megacephalus [5], Makalata sp. [1], Mesomys hispidus [3], Mus musculus [1], Oecomys bicolor [18], Neacomys amoenus [7], Necromys lasiurus [2], Oecomyscleberi [3], Oecomys paricola [4], Oecomys roberi [7], Oecomys aff. Catherinae [1], Oligoryzomys cf. mattrogrossae [5], Oxymycterus amazonicus [1], Proechimys roberti [14], and Proechimys sp. [5]; marsupials: Caluromys philander [6], Cryptonanus sp. [10], Didelphis marsupialis [34], Gliorina venusta [1], Gracilinanus spueranus [5], Marmosa constantiae [64], Marmosa murina [2], Marmosops [13], Marmosops aff. Pinheiroi [5], Metachirus nadicaudatus [5], Monodelphis [4], and Monodelphis saci [2]. Mammals species of Barboza de Oliveira et al. [131]: 51 wild rodents: Thrichomys laurentius [40], Rhipidomys cariri [3], Calomys expulsus [2], Wiedomys pyrrhorhinos [2], Galea spixii [1], Kerodon rupestris [1], and Rattus rattus [2]; 18 marsupials: Monodelphis domestica [9], Didelphis albiventris [7], and Gracilinanus agilis [2]; and 1 wild canid: Cercodyon thous [1]. Foxes species from Di Cataldo [133]: South American grey foxes (Lycalopex griseus) [12] and Andean foxes (L. culpaeus) [9] from the Steppe-Mediterranean region (around Santiago city); Darwin's foxes from the Chiloé Island [10] and one from Temperate Warm Rainy area [11]. *Bats species from Muller et al. [147•]: Myotis chiloensis [27], Histiotus montanus [8], Lasiurus varius [7], Histiotus macrotus [3], L. cinereus [1], and Tadarida brasiliensis [9]. Species from Gardon et al. [107]: rodents: Mus musculus [58], Proechimys sp. (P. cawieri and P. cayennensis) [26], Rattus rattus [17], and other rodents [16]; marsupials: Philander opossum [36], Didelphis marsupialis [4], and other marsupials [2]; chiropters: Molossus molossus [57], Phyllostomushastatus [17], and other marsupials [12]; and birds [83] Prognecnalybea and Prognetapera (swallows from the jail of Rémire-Montjoly, where several human cases were reported (doubtful results)) and batrachians Buffo marinus [21]. Leptodacrylus pentadacylus [20], and other batrachians [6]. Rodents species from Marié et al. 2013 [153]: Micoureus demerarae [1], Mus musculus [3], Oecomys auyantepu [3], Oecomys rutilus [1], Proechimys cayennensis [3], Proechimys cawieri [4], Proechimys sp. [4], Rattus rattus [6], and Rhipidomys nitela [2]. Bats species from Davoust et al. [153]: Pteropitryx macrotis [1], Nocilio albiventris [14], Pteronotus gymnonotus [5], Pteronotus parnellii [37], Pteronotus davyi [1], Anoura caudifera [2], Anoura geoffroyi [15], Glossophaga soricina [2], Phyllostomus elongatus [13], Carollia perspicillata [109], Arthebus planirostris [8], Myotis riparius [1], Eumops auripendulus [9], Molossus barnest [4], and Molossus molossus [11]. Birds species from Destoop, Edouard, and Epelboin: species: chestnut-bellied seed finch (Oryzoborus angolensis) (n=5); domestic canary (Serinus canaria forma domestica) (n=1); ruddy-breasted seedeater (Sporophila minutia) (n=4); and wing-barred seedeater (Sporophila americana) (n=1). Species from Mendoza-Roldan [143]: reptiles: snakes: Chironius multiventris, Chironius scurrulus, Corallus hortulanus, Oxyphrus melanogenys, Philodryas viridissima, Bothrops insularis, Crotalus durissus terrificus, Xenodon neuwiedii, Dipsas turgidus, and Dipsas neuwiedi Bothrops leucurus; lizards: Philodryas nattereri, Hemidactylus mabouia, Salvador merianaes, and Tropidurus catalanensis. Species from Ikeda et al. [144]: bats: Arthebus lituratus, Arthebus planirostris, Myotis nigricans, Platyrhinnus lineatus, Eptesicus furinalis, Carolina perspicillata, Molossus molossus, Phyllostomus discolor, Molossops temminckii, Eumops perotis, and Chiroderma villosum. Mammals species from De Oliveira et al.: Bradypus tridacylus [103], Bradypus sp. [3], Choloepus didacylus [5], Choloepus sp. [31], Tamandua tetradactyla [40], Myrmecophaga tridactyla [67], Cabassous unincicus [4], Dasypus novemcincus [14], Euphractus sexcinctus [14], and Priodontes maximus [25]

<figure><img src="image_22.png" /></figure>
<table><caption>**Table 3** (continued)</caption><thead><tr><th>Author</th><th>Year of the study</th><th>Year of publication</th><th>Language</th><th>Country</th><th>Location</th><th>Methodology</th><th>Sample type</th><th>Species</th><th>Number of samples</th><th>Result</th><th>Comments</th><th>Published</th></tr></thead><tbody><tr><td>Destoop</td><td>2018</td><td></td><td>French</td><td>French Guiana</td><td>Cayenne</td><td>qPCR (IS111)</td><td>Stool</td><td>Captive song-birds + +</td><td>0/11</td><td>0%</td><td>Unpublished data</td><td>No</td></tr><tr><td>Somma-Moreira [115, 116]</td><td>1985</td><td>1987</td><td>English and Spanish</td><td>Uruguay</td><td>NA</td><td>MAT</td><td>Serum</td><td>Birds</td><td>0/66</td><td>0%</td><td></td><td>Yes</td></tr><tr><td>Hernandez [155]</td><td>2003–2004</td><td>2007</td><td>English</td><td>Uruguay</td><td>Dpt of Maldonado</td><td>IFA</td><td>Serum</td><td>Pampas deer (Ozotoceros bezoarticus)</td><td>5/22</td><td>22%</td><td>Experimental wildlife breeding station department</td><td>Yes</td></tr></tbody></table>

CFT, complement fixation test; MA, microagglutination test; ELISA, enzyme-linked immunosorbent assay; IFA, immunofluorescence assay; PCR, qualitative polymerase chain reaction; qPCR, quantitative real-time polymerase chain reaction; MLVA, multilocus variable number tandem repeats analysis; MST, multispacer sequence type.; IHC, immunohistochemistry

*Species from Mares-Guia et al. [77]: Rodentia: Akodon cursor (n=10), Nectomys squamipes [1], Oligoryzomys nigripes [1], Didelphimorpha: Didelphis aurita (n=9); Micoureus (marmosa) paraguayanus (n=12); and Philander frenatus (n=27). Rodents species from Rozenthal et al. [139]: Akodon cursor [32], Mus musculus [19], Oligoryzomys nigripes [16], Delomys doralis [14], Oxymycteris dasytrichus [12], Euryoryzomys russipes [7], Trinomys iheringi [6], Nectomys squamipes [5], Akodon lindberghi [5], Trinomys dimidiatus [4], Akodon montensis [3], Oligoryzomys flavescens [2], Brucepatersonius iheringi [1], Calomys tener [1], Necromys lasiurus [1], Oecomys catherinae [1], Oxymycterus delator [1], and Rhipidomys itoan [1]. Bats species from Ferreira et al. [140]: Carollia perspicillata (n=34), Desmodus rotundus [15], Arthebus lituratus [14], Sturnira lilium [12], Arthebus fimbriatus [7], Rhinophylla pumilio [7] Arthebus planirostris [5], Dermanura cinerea [4], Phyllostomus discolor [4], Arthebus obscurus [2], Glossophaga soricina [2], Myotis nigricans [2], Sturnira tildae [2], Vampyressa pusilla [2], Anoura caudifer [1], Chiroderma doriae [1], Lonchophylla peracchii [1], Micronycteris minuta [1], Micronycteris sp. [1], Phyllostomus hastatus [1], and Trinomycteris nicefori [1]. Free-living deer species from Zanatto et al. [141]: Blastocerus dichotomus [156], Mazama gouazoubira [27], M. bororo [4], M. americana [3], and Ozotocerus bezoarticus [11]. Rodents and marsupials species from Colle [142]: rodents: Euryoryzomys nitidus [1], Hylaeamys megacephalus [5], Makalata sp. [1], Mesomys hispidus [3], Mus musculus [1], Oecomys bicolor [18], Neacomys amoenus [7], Necromys lasiurus [2], Oecomyscleberi [3], Oecomys paricola [4], Oecomys roberi [7], Oecomys aff. Catherinae [1], Oligoryzomys cf. mattrogrossae [5], Oxymycterus amazonicus [1], Proechimys roberti [14], and Proechimys sp. [5]; marsupials: Caluromys philander [6], Cryptonumys sp. [10], Didelphis marsupialis [34], Gliorina venusta [1], Gracilinanus speruanus [5], Marmosa constantiae [64], Marmosa murina [2], Marmosops [13], Marmosops aff. Pinheiroi [5], Metachirus nadicaudatus [5], Monodelphis [4], and Monodelphis saci [2]. Mammals species of Barboza de Oliveira et al. [131]: 51 wild rodents: Thrichomys laurentius [40], Rhipidomys cariri [3], Calomys expulsus [2], Wiedomys pyrrhorhinos [2], Galea spixii [1], Kerodon rupestris [1], and Rattus rattus [2]; 18 marsupials: Monodelphis domestica [9], Didelphis albiventris [7], and Gracilinanus agilis [2]; and 1 wild canid: Cerdocyon thous [1]. Foxes species from Di Cataldo [133]: South American grey foxes (Lycalopex griseus) [12] and Andean foxes (L. culpaeus) [9] from the Steppe-Mediterranean region (around Santiago city); Darwin's foxes from the Chiloé Island [10] and one from Temperate Warm Rainy area [11]. *Bats species from Muller et al. [147•]: Myotis chiloensis [27], Histiotus montanus [8], Lasiurus varius [7], Histiotus macrotus [3], L. cinereus [1], and Tadarida brasiliensis [9]. Species from Gardon et al. [107]: rodents: Mus musculus [58], Proechimys sp. (P. caviери and P. cayennensis) [26], Rattus rattus [17], and other rodents [16]; marsupials: Philander opossum [36], Didelphis marsupialis [4], and other marsupials [2]; chiropters: Molossus molossus [57], Phyllostomushastatus [17], and other marsupials [12]; and birds [83] Prognecnalybea and Prognetapera (swallows from the jail of Rémire-Montjoly, where several human cases were reported (doubtful results)) and batrachians Buffo marinus [21], Leptodacrylus pentadacylus [20], and other batrachians [6]. Rodents species from Marié et al. 2013 [153]: Micoureus demerarae [1], Mus musculus [3], Oecomys auyantepu [3], Oecomys rutilus [1], Proechimys cayennensis [3], Proechimys caviери [4], Proechimys sp. [4], Rattus rattus [6], and Rhipidomys nitela [2]. Bats species from Davoust et al. [153]: Pteropitryx macrotis [1], Nocilio albiventris [14], Pteronotus gymnonotus [5], Pteronotus parnellii [37], Pteronotus davyi [1], Anoura caudifera [2], Anoura geoffroyi [15], Glossophaga soricina [2], Phyllostomus elongatus [13], Carollia perspicillata [109], Arthebus planirostris [8], Myotis riparius [1], Eumops auripendulus [9], Molossus barnest [4], and Molossus molossus [11]. Birds species from Destoop, Edouard, and Epelboin: species: chestnut-bellied seed finch (Oryzoborus angolensis) (n=5); domestic canary (Serinus canaria forma domestica) (n=1); ruddy-breasted seedeater (Sporophila minutia) (n=4); and wing-barred seedeater (Sporophila americana) (n=1). Species from Mendoza-Roldan [143]: reptiles: snakes: Chironius multiventris, Chironius scurrulus, Corallus hortulanus, Oxyphrus melanogenys, Philodyryas viridissima, Bothrops insularis, Crotalus durissus terrificus, Xenodon neuwiedii, Dipsas turgidus, and Dipsas neuwiedi Bothrops leucurus; lizards: Philodyryas nattereri, Hemidactylus mabouia, Salvador merianaes, and Tropidurus catalanensis. Species from Ikeda et al. [144]: bats: Arthebus lituratus, Arthebus planirostris, Myotis nigricans, Platyrhinnus lineatus, Eptesicus furinalis, Carolina perspicillata, Molossus molossus, Phyllostomus discolor, Molossops temminckii, Eumops perotis, and Chiroderma villosum. Mammals species from De Oliveira et al.: Bradypus tridacylus [103], Bradypus sp. [3], Choloepus didacylus [5], Choloepus sp. [31], Tamandua tetradactyla [40], Myrmecophaga tridactyla [67], Cabassous unincicus [4], Dasypus novemcincius [14], Euphractus sexcinctus [14], and Priodontes maximus [25]

<figure><img src="image_22.png" /></figure>
<table><caption>**Table 4** Summary of wildlife species with molecular evidence for *Coxiella burnetii* in Latin America</caption><thead><tr><th>Order/sub-order</th><th>Family</th><th>Species English name</th><th>Species Latin name</th><th>Number</th><th>Year of publication</th><th>Country</th><th>Place</th><th>Publication</th><th>Cycle threshold</th></tr></thead><tbody><tr><td>Pilosa</td><td>Bradypodidae</td><td>3-toed sloth</td><td>Bradypus tridactylus</td><td>1</td><td>2014</td><td>French Guiana</td><td>Cayenne</td><td>Davoust [109]</td><td>23</td></tr><tr><td>Cingulata</td><td>Dasypodidae</td><td>9-banded armadillo</td><td>Dasypus novemcinctus</td><td>1</td><td>2013</td><td>French Guiana</td><td>NA</td><td>Marié [153]</td><td>&gt; 35</td></tr><tr><td>Suina</td><td>Tayassuidae</td><td>White-lipped peccary</td><td>Tayassu pecari</td><td>3</td><td>—</td><td>—</td><td>—</td><td>—</td><td>&gt; 35</td></tr><tr><td>Suina</td><td>Tayassuidae</td><td>Collared peccary</td><td>Pecari tajacu</td><td>3</td><td>—</td><td>—</td><td>—</td><td>—</td><td>&gt; 35</td></tr><tr><td>Perissodac-tyla</td><td>Tapiridae</td><td>South American tapir</td><td>Tapirus terrestris</td><td>2</td><td>—</td><td>—</td><td>—</td><td>—</td><td>1 &gt; 35; 1 &lt; 35</td></tr><tr><td>Rodentia</td><td>Cricetidae</td><td>Cursor grass mouse</td><td>Akodon cursor</td><td>3</td><td>2017</td><td>Brazil</td><td>Rio de Janeiro Atlantic forest</td><td>Rozental [139]</td><td>$</td></tr><tr><td>Rodentia</td><td>Muridae</td><td>House mouse</td><td>Mus musculus</td><td>1</td><td>—</td><td>—</td><td>—</td><td>—</td><td>$</td></tr><tr><td>Rodentia</td><td>Cricetidae</td><td>Atlantic Forest hocicudo</td><td>Oxymycterus dasytrichus</td><td>1</td><td>—</td><td>—</td><td>—</td><td>—</td><td>$</td></tr><tr><td>Rodentia</td><td>Cricetidae</td><td>Black-footed pygmy rice rat</td><td>Oligoryzomys nigripes</td><td>1</td><td>—</td><td>—</td><td>—</td><td>—</td><td>$</td></tr><tr><td>Rodentia</td><td>Echimyidae</td><td>Spiny rat</td><td>Proechimys cuvieri</td><td>1</td><td>2013</td><td>French Guiana</td><td>NA</td><td>Marié [153]</td><td>&gt; 35</td></tr><tr><td>Rodentia</td><td>Caviidae</td><td>Capybara</td><td>Hydrochoerus hydrochaeris</td><td>1</td><td>2019</td><td>French Guiana</td><td>Comté River, Roura</td><td>Christen [154]</td><td>31</td></tr><tr><td>Rodentia</td><td>Caviidae</td><td>Capybara</td><td>Hydrochoerus hydrochaeris</td><td>2</td><td>2013</td><td>French Guiana</td><td>NA</td><td>Marié [153]</td><td>1 &gt; 35; 1 &lt; 35</td></tr><tr><td>Chiroptera</td><td>Phyllostomidae</td><td>Seba's short-tailed bat</td><td>Carollia perspicillata</td><td>3</td><td>—</td><td>—</td><td>Cayenne, Régina, and Saint Jean du Maroni</td><td>—</td><td>&gt; 35</td></tr><tr><td>Chiroptera</td><td>Mormoopidae</td><td>Parnell's mustached bat</td><td>Pteronotus parnellii</td><td>1</td><td>—</td><td>—</td><td>Cayenne, Régina</td><td>—</td><td>&gt; 35</td></tr><tr><td>Chiroptera</td><td>Phyllostomidae</td><td>Lesser spear-nosed bat</td><td>Phyllostomus elongatus</td><td>1</td><td>—</td><td>—</td><td>Régina</td><td>—</td><td>&gt; 35</td></tr><tr><td>Chiroptera</td><td>Noctilionidae</td><td>Lesser bulldog bat</td><td>Noctilio albigentris</td><td>1</td><td>—</td><td>—</td><td>Saint Jean du Maroni</td><td>—</td><td>&gt; 35</td></tr><tr><td>Chiroptera</td><td>Phyllostomidae</td><td>Fringed fruit-eating bat</td><td>Artibeus fimbriatus</td><td>1</td><td>2018</td><td>Brazil</td><td>Atlantic Forest</td><td>Ferreira [140]</td><td>$</td></tr><tr><td>Chiroptera</td><td>Phyllostomidae</td><td>Great fruit-eating bat</td><td>Artibeus lituratus</td><td>3</td><td>—</td><td>—</td><td>—</td><td>—</td><td>$</td></tr><tr><td>Chiroptera</td><td>Molossidae</td><td>Brazilian free-tailed Bats</td><td>Tadarida brasiliensis</td><td>5</td><td>2020</td><td>Chile</td><td>Metropolitan Region</td><td>Muller [147●]</td><td>33.05, 33.55, 34.06, 35.34, and 36.45</td></tr></tbody></table>
<table><caption>**Table 4** Summary of wildlife species with molecular evidence for *Coxiella burnetii* in Latin America</caption><thead><tr><th>Order/sub-order</th><th>Family</th><th>Species English name</th><th>Species Latin name</th><th>Number</th><th>Year of publication</th><th>Country</th><th>Place</th><th>Publication</th><th>Cycle threshold</th></tr></thead><tbody><tr><td>Pilosa</td><td>Bradypodidae</td><td>3-toed sloth</td><td>Bradypus tridactylus</td><td>1</td><td>2014</td><td>French Guiana</td><td>Cayenne</td><td>Davoust [109]</td><td>23</td></tr><tr><td>Cingulata</td><td>Dasypodidae</td><td>9-banded armadillo</td><td>Dasypus novemcinctus</td><td>1</td><td>2013</td><td>French Guiana</td><td>NA</td><td>Marié [153]</td><td>&gt; 35</td></tr><tr><td>Suina</td><td>Tayassuidae</td><td>White-lipped peccary</td><td>Tayassu pecari</td><td>3</td><td>—</td><td>—</td><td>—</td><td>—</td><td>&gt; 35</td></tr><tr><td>Suina</td><td>Tayassuidae</td><td>Collared peccary</td><td>Pecari tajacu</td><td>3</td><td>—</td><td>—</td><td>—</td><td>—</td><td>&gt; 35</td></tr><tr><td>Perissodac-tyla</td><td>Tapiridae</td><td>South American tapir</td><td>Tapirus terrestris</td><td>2</td><td>—</td><td>—</td><td>—</td><td>—</td><td>1 &gt; 35; 1 &lt; 35</td></tr><tr><td>Rodentia</td><td>Cricetidae</td><td>Cursor grass mouse</td><td>Akodon cursor</td><td>3</td><td>2017</td><td>Brazil</td><td>Rio de Janeiro Atlantic forest</td><td>Rozental [139]</td><td>$</td></tr><tr><td>Rodentia</td><td>Muridae</td><td>House mouse</td><td>Mus musculus</td><td>1</td><td>—</td><td>—</td><td>—</td><td>—</td><td>$</td></tr><tr><td>Rodentia</td><td>Cricetidae</td><td>Atlantic Forest hocicudo</td><td>Oxymycterus dasytrichus</td><td>1</td><td>—</td><td>—</td><td>—</td><td>—</td><td>$</td></tr><tr><td>Rodentia</td><td>Cricetidae</td><td>Black-footed pygmy rice rat</td><td>Oligoryzomys nigripes</td><td>1</td><td>—</td><td>—</td><td>—</td><td>—</td><td>$</td></tr><tr><td>Rodentia</td><td>Echimyidae</td><td>Spiny rat</td><td>Proechimys cuvieri</td><td>1</td><td>2013</td><td>French Guiana</td><td>NA</td><td>Marié [153]</td><td>&gt; 35</td></tr><tr><td>Rodentia</td><td>Caviidae</td><td>Capybara</td><td>Hydrochoerus hydrochaeris</td><td>1</td><td>2019</td><td>French Guiana</td><td>Comté River, Roura</td><td>Christen [154]</td><td>31</td></tr><tr><td>Rodentia</td><td>Caviidae</td><td>Capybara</td><td>Hydrochoerus hydrochaeris</td><td>2</td><td>2013</td><td>French Guiana</td><td>NA</td><td>Marié [153]</td><td>1 &gt; 35; 1 &lt; 35</td></tr><tr><td>Chiroptera</td><td>Phyllostomidae</td><td>Seba's short-tailed bat</td><td>Carollia perspicillata</td><td>3</td><td>—</td><td>—</td><td>Cayenne, Régina, and Saint Jean du Maroni</td><td>—</td><td>&gt; 35</td></tr><tr><td>Chiroptera</td><td>Mormoopidae</td><td>Parnell's mustached bat</td><td>Pteronotus parnellii</td><td>1</td><td>—</td><td>—</td><td>Cayenne, Régina</td><td>—</td><td>&gt; 35</td></tr><tr><td>Chiroptera</td><td>Phyllostomidae</td><td>Lesser spear-nosed bat</td><td>Phyllostomus elongatus</td><td>1</td><td>—</td><td>—</td><td>Régina</td><td>—</td><td>&gt; 35</td></tr><tr><td>Chiroptera</td><td>Noctilionidae</td><td>Lesser bulldog bat</td><td>Noctilio albigentris</td><td>1</td><td>—</td><td>—</td><td>Saint Jean du Maroni</td><td>—</td><td>&gt; 35</td></tr><tr><td>Chiroptera</td><td>Phyllostomidae</td><td>Fringed fruit-eating bat</td><td>Artibeus fimbriatus</td><td>1</td><td>2018</td><td>Brazil</td><td>Atlantic Forest</td><td>Ferreira [140]</td><td>$</td></tr><tr><td>Chiroptera</td><td>Phyllostomidae</td><td>Great fruit-eating bat</td><td>Artibeus lituratus</td><td>3</td><td>—</td><td>—</td><td>—</td><td>—</td><td>$</td></tr><tr><td>Chiroptera</td><td>Molossidae</td><td>Brazilian free-tailed Bats</td><td>Tadarida brasiliensis</td><td>5</td><td>2020</td><td>Chile</td><td>Metropolitan Region</td><td>Muller [147●]</td><td>33.05, 33.55, 34.06, 35.34, and 36.45</td></tr></tbody></table>
<table><caption>**Table 4** (continued)</caption><thead><tr><th>Order/sub-order</th><th>Family</th><th>Species English name</th><th>Species Latin name</th><th>Number</th><th>Year of publication</th><th>Country</th><th>Place</th><th>Publication</th><th>Cycle threshold</th></tr></thead><tbody><tr><td>Chiroptera</td><td>Phyllostomidae</td><td>Seba's short-tailed bat</td><td>Carollia spicillata</td><td>3</td><td>2022</td><td>Colombia</td><td>Macaragua cave, located in Las Vueltas village, Municipality of Curiti, Santander Department</td><td>Silva-Ramos [150•]</td><td>NA</td></tr><tr><td>Chiroptera</td><td>Mormoopidae</td><td>Ghost-faced bat</td><td>Mormops megalophylla</td><td>2</td><td>—</td><td>—</td><td>—</td><td>—</td><td>NA</td></tr><tr><td>Chiroptera</td><td>Natalidae</td><td>Trinidadian funnel-eared bat</td><td>Natalus tumidirostris</td><td>3</td><td>—</td><td>—</td><td>—</td><td>—</td><td>NA</td></tr></tbody></table>

NA, data not available

$, In this work, we used “conventional” PCR followed by electrophoreses, which is why we will not be able to provide a CT value

Among Latin American countries, Brazil has the largest number of publications, with evidence of *C. burnetii* circulation in livestock, pets, wild animals, and ticks. Bovines are the species with the highest average, followed by sheep, goats, dogs, wild cervids, wild rodents, bats, and ticks. This scenario is probably linked to the economic importance of the sector since Brazil is the second-largest producer of beef cattle and has a large dairy herd. Three papers are related to the detection of *C. burnetii* in animals with reproductive disorders [79, 82, 88••]. The study conducted in the state of Alagoas, in the northeast of Brazil, described a goat herd with high frequency in serology (55.1%) and molecular detection by conventional PCR in two placental samples from seropositive animals (8.7%) [79]. Another survey on cattle aborted fetuses reported 9.2% (7/76) of positivity at qPCR, with 3 positive fetuses from the state of Rio Grande do Sul and 4 positive fetuses from the state of São Paulo, reinforcing the widespread distribution of the pathogen in the country [88••]. In both studies, however, the lack of histopathological assay does not confirm that *C. burnetii* was responsible for the abortions [200]. The vast majority of studies in Brazil are based on serological and molecular detection in convenience samples. Only recently, statistical design studies are being carried out to understand better *C. burnetii* epidemiology [64, 79–81, 140, 141]. In the state of São Paulo, a study with 1515 cattle samples from 54 cities collected in 9 different slaughterhouses reported a seroprevalence of 23.8% using an in-house IFA and the detection of *C. burnetii* by qPCR in 12.2% of the seropositive animals, highlighting the risk of infection for abattoir workers [64].

Molecular data suggest the existence of diverse MST and MLVA genotypes of *C. burnetii* in the state of São Paulo (ST74) [86]. In the publication, an ST73 was also detected in samples from Argentina. These genotypes were only reported in South America, reinforcing the need for epidemiological studies aiming at the isolation and characterization of human and animal strains to better understand the real impact on animal and public health [86].

Based on a recent analysis of typing methods and their correlation with the genomic groups [201], cattle and goat strains from Brazil belong to a genomic group (GG)-III, ovine strains from Brazil are correlated with GG-I, the same as ST17 detected in French Guiana, and the Argentinian strain At12 is placed in GG-IVb-associated, a genomic group with genetic relation with the Australian strains AuQ01, AuQ02, and Namibia (ST-30), which are isolates placed on GG-X, and strains placed in the GG-IV, like the isolate Leningrad-2 (ST-7). Strains from GG-III are usually detected in cattle and are rarely involved in human infection [201, 202]. On the other hand, *C. burnetii* belonging to GG-I can be found across the globe and shows elevated virulence, and strains from GG-IVb and GG-X were mainly isolated from human cases, illustrating the virulence potential of Brazilian ovine strains and the Argentinian isolate At12 described by Pacheco et al. [201, 203].

Although the transmission of *C. burnetii* through the consumption of contaminated milk from dairy animals remains controversial, studies conducted in the last decade in the state of Minas Gerais have shown samples of artisanal Minas cheese contaminated by this proteobacterium [79, 85, 87]. The risk has been assessed; in some cases, the consumption
<table><caption>**Table 4** (continued)</caption><thead><tr><th>Order/sub-order</th><th>Family</th><th>Species English name</th><th>Species Latin name</th><th>Number</th><th>Year of publication</th><th>Country</th><th>Place</th><th>Publication</th><th>Cycle threshold</th></tr></thead><tbody><tr><td>Chiroptera</td><td>Phyllostomidae</td><td>Seba's short-tailed bat</td><td>Carollia spicillata</td><td>3</td><td>2022</td><td>Colombia</td><td>Macaragua cave, located in Las Vueltas village, Municipality of Curiti, Santander Department</td><td>Silva-Ramos [150•]</td><td>NA</td></tr><tr><td>Chiroptera</td><td>Mormoopidae</td><td>Ghost-faced bat</td><td>Mormops megalophylla</td><td>2</td><td>—</td><td>—</td><td>—</td><td>—</td><td>NA</td></tr><tr><td>Chiroptera</td><td>Natalidae</td><td>Trinidadian funnel-eared bat</td><td>Natalus tumidirostris</td><td>3</td><td>—</td><td>—</td><td>—</td><td>—</td><td>NA</td></tr></tbody></table>

NA, data not available

$, In this work, we used “conventional” PCR followed by electrophoreses, which is why we will not be able to provide a CT value

Among Latin American countries, Brazil has the largest number of publications, with evidence of *C. burnetii* circulation in livestock, pets, wild animals, and ticks. Bovines are the species with the highest average, followed by sheep, goats, dogs, wild cervids, wild rodents, bats, and ticks. This scenario is probably linked to the economic importance of the sector since Brazil is the second-largest producer of beef cattle and has a large dairy herd. Three papers are related to the detection of *C. burnetii* in animals with reproductive disorders [79, 82, 88••]. The study conducted in the state of Alagoas, in the northeast of Brazil, described a goat herd with high frequency in serology (55.1%) and molecular detection by conventional PCR in two placental samples from seropositive animals (8.7%) [79]. Another survey on cattle aborted fetuses reported 9.2% (7/76) of positivity at qPCR, with 3 positive fetuses from the state of Rio Grande do Sul and 4 positive fetuses from the state of São Paulo, reinforcing the widespread distribution of the pathogen in the country [88••]. In both studies, however, the lack of histopathological assay does not confirm that *C. burnetii* was responsible for the abortions [200]. The vast majority of studies in Brazil are based on serological and molecular detection in convenience samples. Only recently, statistical design studies are being carried out to understand better *C. burnetii* epidemiology [64, 79–81, 140, 141]. In the state of São Paulo, a study with 1515 cattle samples from 54 cities collected in 9 different slaughterhouses reported a seroprevalence of 23.8% using an in-house IFA and the detection of *C. burnetii* by qPCR in 12.2% of the seropositive animals, highlighting the risk of infection for abattoir workers [64].

Molecular data suggest the existence of diverse MST and MLVA genotypes of *C. burnetii* in the state of São Paulo (ST74) [86]. In the publication, an ST73 was also detected in samples from Argentina. These genotypes were only reported in South America, reinforcing the need for epidemiological studies aiming at the isolation and characterization of human and animal strains to better understand the real impact on animal and public health [86].

Based on a recent analysis of typing methods and their correlation with the genomic groups [201], cattle and goat strains from Brazil belong to a genomic group (GG)-III, ovine strains from Brazil are correlated with GG-I, the same as ST17 detected in French Guiana, and the Argentinian strain At12 is placed in GG-IVb-associated, a genomic group with genetic relation with the Australian strains AuQ01, AuQ02, and Namibia (ST-30), which are isolates placed on GG-X, and strains placed in the GG-IV, like the isolate Leningrad-2 (ST-7). Strains from GG-III are usually detected in cattle and are rarely involved in human infection [201, 202]. On the other hand, *C. burnetii* belonging to GG-I can be found across the globe and shows elevated virulence, and strains from GG-IVb and GG-X were mainly isolated from human cases, illustrating the virulence potential of Brazilian ovine strains and the Argentinian isolate At12 described by Pacheco et al. [201, 203].

Although the transmission of *C. burnetii* through the consumption of contaminated milk from dairy animals remains controversial, studies conducted in the last decade in the state of Minas Gerais have shown samples of artisanal Minas cheese contaminated by this proteobacterium [79, 85, 87]. The risk has been assessed; in some cases, the consumption

<figure><img src="image_23.png" /></figure>
<figure><img src="image_23.png" /><figcaption>**Fig. 5** Map of the publications on *Coxiella burnetii* infection in ticks in Latin America and the Caribbean</figcaption></figure>

of dairy products from infected animals may induce seroconversion but no clinical manifestation [204].

In conclusion, *C. burnetii* infection in Brazil has been reported in different domestic and wild animal species. Although the pathogen seems widespread in the country, studies are concentrated in only six states. Hence, the prevalence of *C. burnetii* in animals is spatially biased. More studies are needed, especially in states without data, to better understand Q fever epidemiology in Brazil and its implications for animal and human health. Increasing Q fever knowledge would benefit preventive and control measures, reducing the risk of human infection and economic losses in livestock.

## Argentina

*Coxiella burnetii* is a pathogen understudied in Argentina, with few or no publications in the second half of the twentieth century and early twenty-first. Serological studies have covered farm animals, pets, and humans, although the diversity of techniques and cutting-edge titles used make their comparison difficult. Only two publications use diagnosis through molecular biology, and they are also the only ones that studied ticks. Q fever is currently subject to compulsory notification.

The description of the first human cases of Q fever by serology in Argentina was carried out in 1957 in Córdoba

[205] and in 1959 by a truck driver traveling through the provinces of Chaco, Formosa, Corrientes, and Missions [206]. The first human case in Buenos Aires City (BAC) was reported in 1962 (Ruggiero et al. 1962, unpublished data). Numerous serological studies were carried out in the 1950s and 1960s on different animal species, finding different seropositivities (Table 1) [68–71].

Notably, after these pioneering studies, there were no publications on the subject until 1997, when the suspicion of Q fever was reported in goats of Entre Ríos (although animals imported from Uruguay) from compatible signs by serology and was also detected in people related to the farm [207, 208]. In 1999, *C. burnetii* was detected in two imported cattle that were in quarantine and were then euthanized [208, 209].

Recently, clinically healthy dogs from poor neighborhoods of BAC were seropositive by indirect immunofluorescence [129], but not by IFI in cats [124, 125, 203, 205] or by molecular biology techniques in *Rhipicephalus sanguineus* sensu lato ticks [210]. Also, in BAC, 1/99 human sera were positive by indirect immunofluorescence [211].

In samples collected in 2004, Trezeguet et al. carried out a serological test using ELISA (phases I and II) as a screening test and FC as a confirmatory test in 840 goats of 56 farms covering practically the whole country; although only 9/840 (1,1%) were positive through FC, all belonged to the
<figure><img src="image_24.png" /><figcaption>**Fig. 5** Map of the publications on *Coxiella burnetii* infection in ticks in Latin America and the Caribbean</figcaption></figure>

of dairy products from infected animals may induce seroconversion but no clinical manifestation [204].

In conclusion, *C. burnetii* infection in Brazil has been reported in different domestic and wild animal species. Although the pathogen seems widespread in the country, studies are concentrated in only six states. Hence, the prevalence of *C. burnetii* in animals is spatially biased. More studies are needed, especially in states without data, to better understand Q fever epidemiology in Brazil and its implications for animal and human health. Increasing Q fever knowledge would benefit preventive and control measures, reducing the risk of human infection and economic losses in livestock.

## Argentina

*Coxiella burnetii* is a pathogen understudied in Argentina, with few or no publications in the second half of the twentieth century and early twenty-first. Serological studies have covered farm animals, pets, and humans, although the diversity of techniques and cutting-edge titles used make their comparison difficult. Only two publications use diagnosis through molecular biology, and they are also the only ones that studied ticks. Q fever is currently subject to compulsory notification.

The description of the first human cases of Q fever by serology in Argentina was carried out in 1957 in Córdoba

[205] and in 1959 by a truck driver traveling through the provinces of Chaco, Formosa, Corrientes, and Missions [206]. The first human case in Buenos Aires City (BAC) was reported in 1962 (Ruggiero et al. 1962, unpublished data). Numerous serological studies were carried out in the 1950s and 1960s on different animal species, finding different seropositivities (Table 1) [68–71].

Notably, after these pioneering studies, there were no publications on the subject until 1997, when the suspicion of Q fever was reported in goats of Entre Ríos (although animals imported from Uruguay) from compatible signs by serology and was also detected in people related to the farm [207, 208]. In 1999, *C. burnetii* was detected in two imported cattle that were in quarantine and were then euthanized [208, 209].

Recently, clinically healthy dogs from poor neighborhoods of BAC were seropositive by indirect immunofluorescence [129], but not by IFI in cats [124, 125, 203, 205] or by molecular biology techniques in *Rhipicephalus sanguineus* sensu lato ticks [210]. Also, in BAC, 1/99 human sera were positive by indirect immunofluorescence [211].

In samples collected in 2004, Trezeguet et al. carried out a serological test using ELISA (phases I and II) as a screening test and FC as a confirmatory test in 840 goats of 56 farms covering practically the whole country; although only 9/840 (1,1%) were positive through FC, all belonged to the