THERYA NOTES 2024, Vol. 5 : 139-149 DOI: 10.12933/therya_notes-24-162 ISSN 2954-3614

The rodent Abrothrix olivacea and its role as a host of micro- and macroparasites in Chile: A systematic review and meta-analysis

El roedor Abrothrix olivacea y su papel como hospedero de micro y macroparásitos en Chile: Una revisión sistemática y metaanálisis

Gabriela Gaona1, Hugo Henríquez1,2, and André V. Rubio1*

1Departamento de Ciencias Biológicas Animales, Facultad de Ciencias Veterinarias y Pecuarias, Universidad de Chile. Av. Santa Rosa 11735, C. P. 8820000. Santiago, Chile. E-mail: g.gaonamarcel@gmail.com (G-G); hugohenriquezf92@gmail.com (H-H); arubio@uchile.cl (AV-R).

2Programa de Doctorado en Ciencias Silvoagropecuarias y Veterinarias, Universidad de Chile. Campus Sur, Av. Santa Rosa 11315, C. P. 8820000. Santiago, Chile.

*Corresponding author

The olive grass mouse (Abrothrix olivacea) is a South American rodent with ecological and life history characteristics that suggest a high reservoir capacity for zoonotic pathogens. The objectives of this study were to determine the micro- and macroparasites of the olive grass mouse in Chile and to compare the prevalence of these parasites in other sympatric rodents. A systematic review was carried out on the micro- and macroparasites of the olive grass mouse in Chile reported in articles available in the PubMed, Web of Science, and Scielo Scientific Library databases using the PRISMA method. In addition, a meta-analysis of the prevalence of these micro- and macroparasites in the olive grass mouse was performed, and their prevalence in other rodents was compared using meta-regression. Of the 51 articles available in the literature, a total of 68 micro- and macroparasites were recorded, of which 6 are zoonotic. Prevalence varied between parasites, highlighting a higher overall prevalence of helminths in the olive grass mouse (8.4 %) compared to other rodents (2.8 %). The presence of micro- and macroparasites and their high prevalence (in some cases, compared to the prevalence observed in other rodents) highlight the potential role of the olive grass mouse in the epidemiology of pathogens. Therefore, further research on disease ecology focused on this rodent species is required.

Key words: Cricetidae; pathogens; small mammals; Rodentia; zoonosis.

El ratón oliváceo (Abrothrix olivacea) es un roedor sudamericano que presenta características ecológicas y de historia de vida que sugieren una alta capacidad reservoria para patógenos zoonóticos. El objetivo de este estudio fue realizar un análisis de los microparásitos y macroparásitos en el ratón oliváceo en Chile, así como comparar la prevalencia con otros roedores simpátricos. Se realizó una revisión sistemática sobre microparásitos y macroparásitos en el ratón oliváceo en Chile consultando las bases de datos PubMed, Web of Science y Scielo Scientific Library, utilizando el método PRISMA. Además, se realizó un metaanálisis de la prevalencia de estos microparásitos y macroparásitos en el ratón oliváceo y se comparó con la prevalencia en otros roedores utilizando metarregresión. De 51 artículos disponibles en la literatura, se registraron 68 micro y macroparásitos en total, de los cuales 6 son zoonóticos. La prevalencia varió entre parásitos, destacando una mayor prevalencia general de helmintos en el ratón oliváceo (8.4 %), en comparación a otros roedores (2.8 %). La presencia de micro y macroparásitos y alta prevalencia (en algunos casos comparado a otros roedores), resalta el potencial papel del ratón oliváceo en la epidemiología de patógenos, por lo que se sugiere una mayor investigación sobre ecología de enfermedades enfocada en esta especie de roedor.

Palabras clave: Cricetidae; patógenos; pequeños mamíferos; Rodentia; zoonosis.

© 2024 Asociación Mexicana de Mastozoología, www.mastozoologiamexicana.org

The order Rodentia is the group of mammals that hosts the largest number of host species of zoonotic pathogens, associated with more than 80 diseases that affect humans (Han et al. 2016). Rodent species considered reservoirs of zoonotic pathogens share several attributes related to their ecology and life history. For instance, they tend to have a wide geographic distribution, are generalists regarding habitat and diet, and exhibit fast life history traits (e.g., rapid growth, early maturity, large litter size; Han et al. 2015; Plourde et al. 2017). Considering these characteristics, recent studies include the olive grass mouse (Abrothrix olivacea, Cricetidae: Abrotrichini) within the group of species with a high capacity to host pathogens with zoonotic potential (Han et al. 2015), above that of many other Neotropical species (Han et al. 2020). In addition, it stands out as a host of possible new Orthohantaviruses (Mull et al. 2022).

The olive grass mouse is a small rodent (30.4 g ± 4.3 g) native to Chile and Argentina. In Chile, this rodent is widely distributed, from Tarapacá to Tierra del Fuego and from sea level to 2,500 m (Spotorno et al. 2000). It inhabits various environments and is highly adapted to anthropic habitats (Iriarte 2008; Moreno-Salas et al. 2020; Barceló et al. 2021). This species has been described as a host of various potentially zoonotic or zoonotic pathogens, such as Leptospira spp., Yersinia enterocolitica, and Cryptosporidium spp. (Llanos-Soto and González-Acuña 2019; Infante et al. 2022).

In this context, aiming to know the current status of the parasites recorded in the olive grass mouse in Chile, this study conducted a systematic review of the microparasites (bacteria, viruses, protozoa, and fungi) and macroparasites (helminths and arthropods) in this rodent. In addition, a meta-analysis of the prevalence of micro- and macroparasites was carried out to record and compare this prevalence with the one reported in other sympatric rodents. This information will give us a general and up-to-date overview of this rodent from an epidemiological standpoint.

A systematic review of scientific articles on micro- and macroparasites of the olive grass mouse in Chile was carried out, guided by the PRISMA statement (preferred reporting items for systematic reviews and meta-analyses; Moher et al. 2014). The survey was carried out in the PubMed, Web of Science (WoS), and Scielo Scientific Library databases using the following Boolean operators and keywords: ((rodentia) OR (rodent*)) OR (rodents)) AND ((pathogen*) OR (bacteria*) OR (helminth*) OR (fungi*) OR (protozoa*) OR (virus*) OR (parasite*) OR (flea*) OR (tick*) OR (mite*) OR (lice*) OR (zoonoses)) AND (Chile)). The articles were compiled according to title and abstract, regardless of language or year of publication, to obtain as much literature as possible. The inclusion criteria were as follows. The studies were carried out in Chile and were related to the detection of micro- or macroparasites in the olive grass mouse. Review articles and duplicates between the different databases were eliminated, and a full-text review was carried out, discarding those publications that did not meet the inclusion criteria. In addition, we performed a manual review and selection of articles that were not identified in the database search as such but were cited in the bibliography of the articles identified.

The information in the articles was extracted and organized in an Excel spreadsheet with the following data: type of parasite, detection of parasites in other rodent species, diagnostic method, author, and year. For the taxonomy of micro- and macroparasites, the names indicated in each article were maintained. Those publications that did not provide quantitative data were included only for the systematic review. On the other hand, those that reported prevalence and sample size were also used to perform a meta-analysis. To this end, the data were again organized in an Excel spreadsheet according to the type of parasite, host species, sample size, number of positive samples, prevalence, author, and year.

For the prevalence analysis, a meta-analysis was performed with random models weighted by sample size, with a 95 % confidence interval (Nikolakopoulou et al. 2014). Heterogeneity between studies was measured with the I2 index (Higgins et al. 2003), and plots were constructed showing the prevalences arranged according to the group of micro- or macroparasites studied, considering 2 subgroups: A. olivacea and "other rodents", the latter considering any other rodent species (native or introduced species) analyzed exclusively in the same studies that included the olive grass mouse. Parasite prevalences according to host were compared using meta-regressions (Baker et al. 2009) carried out according to the groups and subgroups used in the meta-analyses. In addition, within a group of parasites (e.g., helminths), meta-regression analyses were separated when different diagnostic methods were used (e.g., microscopy and PCR analysis). The entire meta-analysis was performed with the the R packages 'metafor' (Viechtbauer 2010) and 'meta' (Balduzzi et al. 2019).

According to the literature search and review, 51 articles were included in the systematic review, and 38 articles were used in the meta-analysis. All included articles are cited in Table 1. In addition, some articles included in the meta-analysis where the prevalence in the olive grass mouse was 0 are listed in Appendix 1. The literature review process is summarized in the flowchart in Appendix 2. The studies are distributed in all administrative regions of Chile, mainly concentrated in the Coquimbo region (north-central zone, with 14 studies, 27.5 %), followed by the Metropolitan region (central zone, 11 studies, 21.6 %) and the Los Ríos region (southern zone, 11 studies, 21.6 %; Figure 1). According to the type of parasite, 13 (25.5 %) articles analyzed ectoparasites, 12 (23.6 %) protozoa, 10 (19.6 %) bacteria, 9 (17.6 %) viruses, and 7 (13.7 %) helminths. No articles were found on parasitic fungi in the olive grass mouse. The largest number of publications (n = 8) reported the Hantavirus (Andes virus, ANDV), the protozoan Trypanosoma cruzi (n = 7), and the bacterium Leptospira sp. (n = 6; Table 1). In total, 68 micro- and macroparasites were recorded, 40 (58.8 %) described at the species level and 28 (41.2 %) to genus, family, or phylum (Table 1). Of the 68 parasites, 52 are macroparasites (35 arthropods and 17 helminths), and 16 are microparasites (8 protozoa, 6 bacteria, and 2 viruses); of the total, 6 are recognized as zoonotic.

First, the analysis of the ANDV virus was performed, as it was the only virus with a sufficient number of publications for the meta-analysis. For this analysis, the subgroups were (1) A. olivacea, (2) Oligoryzomys longicaudatus (main ANDV reservoir), and (3) other rodents. The prevalence of ANDV showed a high heterogeneity between articles (I2 = 92 %). Furthermore, the overall prevalence was higher in O. longicaudatus (4.6 %) compared to the olive grass mouse (0.4 %) and all other rodents (0.9 %; Table 2, Appendix 3). These differences in prevalence among rodents were statistically significant (P < 0.001; Table 3).

For bacteria, 9 studies were included in the meta-analysis. The prevalence of bacteria was highly heterogeneous between studies (I2 = 95 %), showing marked differences in the prevalence of Mycoplasma spp. between the olive grass mouse (75 %) and all other rodents (19.7 %; Appendix 4). In general, there were no significant differences in the prevalence of bacteria in the olive grass mouse (32 %) compared to the other rodents (22 %; P = 0.336; Table 3). Similarly, there were no statistical differences when only the prevalences of Leptospira spp. were compared (P = 0.611; Table 3).

The prevalence of protozoans was also highly heterogeneous between studies (I2 = 97 %, Appendix 5). No differences (P = 0.659) were observed in the prevalence of these microparasites between the olive grass mouse (25.7 %) and other rodents (22.9 %). The protozoa Sarcocystis sp. and members of the subfamily Toxoplasmatinae were found only in the olive grass mouse and not in other rodents. In contrast, Babesia sp. was reported in other rodents but not in the olive grass mouse (Appendix 6). For Trypanosoma cruzi, there was a larger number of publications (n = 7), so a meta-regression was performed for this protozoan, but no significant differences (P = 0.578) were observed between the prevalences of both subgroups (Table 3).

The prevalence of ectoparasites between publications was highly heterogeneous (I2 = 95 %, Appendix 6), and the prevalence in olive mice (10.8 %) was lower than that in other rodents (14.6 %). However, these differences are not statistically significant (P = 0.465; Table 3).

For the analysis of helminths, they were separated into 2 groups according to the diagnostic methodology: 1) studies where helminths were identified by stool examination (ova) and 2) studies where helminths were identified with macroscopic specimens (adults or larvae) after the necropsy of the host (Table 1). The heterogeneity in the prevalence of helminths between publications was high in stool tests (I2 = 87 %) and macroscopic specimens (I2 = 95 %, Table 2). The average prevalence of helminths from stool analyses was significantly different (P = 0.048) between the olive grass mouse (4.5 %) and all other rodents (1.3 %). Regarding the prevalences using data from macroscopic specimens, there was also a higher prevalence in the olive grass mouse (14.6 %) compared to other rodents (5.2 %), although it was not significant (P = 0.066; Table 3; Appendix 7).

In this systematic review, we compiled the published information on macro- and microparasites found in olive grass mice inhabiting different locations in Chile. From a public health approach, the olive grass mouse can host at least 6 pathogens considered zoonotic, such as Leptospira spp., Yersinia enterocolitica, and Orthohantavirus. Pathogenic leptospires have been found mainly in central and southern Chile, and some studies have reported a higher prevalence in olive grass mice than in other rodents (Luna et al. 2020). The fact that we did not find significant differences between prevalences in our meta-analysis makes epidemiological sense because pathogenic Leptospira spp. are generally considered multihost (Boey et al. 2019). However, this type of analysis, which mainly included molecular analyses, is limited by not including the different types of serovars that circulate in rodents. This restricts a more in-depth analysis of the possible differences in the prevalence of Leptospira spp. Concerning Yersinia enterocolitica, only one descriptive study reports this pathogen in different species of Chilean rodents (Zamora et al. 1998) without particularly highlighting the role of the olive grass mouse. In the case of Orthohantavirus, the Andes strain (ANDV) thrives in the geographic range of the olive grass mouse; this strain is considered one of the most important Orthohantaviruses causing the Hantavirus Cardiopulmonary Syndrome (Kruger et al. 2015). For ANDV, the main reservoir is the long-tailed mouse (O. longicaudatus), whereas other rodent species, including the olive grass mouse, have shown antibodies against ANDV. However, it is still unknown whether these rodent species can act as vectors of ANDV to humans; therefore, additional virological studies are needed in these rodent species (Torres-Pérez et al. 2016). According to meta-analyses and other studies of ANDV in rodent communities in Chile and Argentina, the seroprevalence of ANDV tends to be higher in O. longicaudatus compared to other sympatric native rodents (Polop et al. 2010; Rubio et al. 2019; Torres-Pérez et al. 2019). ANDV infection in the olive grass mouse and other native rodents has been suggested to be mainly due to transmission from the main reservoir to these rodents (spillover; Polop et al. 2010; Rubio et al. 2019). However, all studies on Orthohantavirus, including those in this review, assessed only the presence of antibodies and not infection as such. On the other hand, a recent study that carried out predictive models using ecological and phylogenetic information of rodents indicates that the olive grass mouse and other rodents from America could be new hosts of Orthohantavirus (Mull et al. 2022). This highlights the need to conduct further studies on these viruses in the olive grass mouse throughout its range, where other undescribed Orthohantaviruses could exist (Mull et al. 2022). On the other hand, our systematic review evidenced the lack of knowledge about other viruses in the olive grass mouse since the review only identified several studies on hantavirus and a single study on morbillivirus.

This systematic review found other potentially zoonotic biological agents, such as Bartonella spp., Giardia spp. and Cryptosporidium spp., but since they have been identified at the genus level, there is no certainty that they are zoonotic species. In Chile, there are cases of giardidiosis and cryptospyridosis in humans within the geographic range of the olive grass mouse (Neira-Otero et al. 2005; Vidal et al. 2010). Therefore, further studies are required to deepen the taxonomic knowledge of these biological agents.

Ticks, mites, and fleas have been described in the olive grass mouse to date (Table 1). These ectoparasites may be important vectors of zoonoses. An interesting case to highlight is scrub typhus fever, an emerging disease whose causative agent is a rickettsia (Orientia spp.) and in which the olive grass mouse could play an important role. This vectorial disease (transmitted by thrombiculid mites) has recently been described in Chile (Abarca et al. 2020; Weitzel et al. 2021), highlighting a high infestation of thrombiculids infested with this rickettsia in the olive grass mouse (Acosta-Jamett et al. 2020; Silva-de la Fuente et al. 2023). Since the olive grass mouse is one of the native species of Chile that is most adapted to anthropic settlements (Moreno-Salas et al. 2020; Barceló et al. 2021), this rodent species could increase the transmission of scrub typhus fever by increasing the likelihood of contact between vector mites and humans.

Regarding fleas, some species parasitizing the olive grass mouse have been reported (Hectopsylla spp., Neotyphloceras chilensis, Neotyphloceras pardinasi, Sphinctopsylla ares, Tetrapsyllus rhombus, and Nosopsyllus fasciatus), which have previously been recognized as carriers of Bartonella spp. in Rattus rattus, Abrothrix sp., and other rodents in Chile (Moreno-Salas et al. 2019; Müller et al. 2020). In addition, 2 studies highlight the high richness of these ectoparasites in the olive grass mouse (Bazán-León et al. 2013; Moreno-Salas et al. 2020). Moreno-Salas et al. (2020) emphasize a higher richness of flea species in the olive grass mouse compared to all the other rodent species analyzed, as well as a greater number of rickettsia-positive fleas. Molecular analyses of these rickettsias indicated that the sequences are close to the rickettsia of the spotted fever group, so there is a zoonotic potential not yet studied in Chile (Moreno-Salas et al. 2020). Descriptions of ticks in the olive grass mouse are scarce since only the species Ixodes abrocomae (Guglielmone et al. 2010) and a larva of Ixodes sp. (Muñoz-Leal et al. 2019) have been described.

Overall, there is a larger number of zoonotic helminths described in rodents compared to other zoonotic biological agents such as viruses, bacteria, and protozoa (Han et al. 2016). In Chile, all the studies in this review (Table 1) have described helminths in the olive grass mouse by macro- and microscopic analyses, mainly at the family or genus levels. This makes it difficult to identify whether there are zoonotic helminth species in the olive grass mouse, although many genera of helminths described in this rodent species infect humans (i.e., Moniliformis, Syphacia, Capillaria, Trichuris, and Hymenolepis; Salehabadi et al. 2008; Gomes da Rocha et al. 2015; Panti-May et al. 2020). The high prevalence (greater than 20 %) of certain helminths in the olive grass mouse (Physaloptera calnuensis, Syphacia sp., Trichuris sp., Pterigodermatites sp. Hymenopepis sp., and Protospirura sp.; Landaeta-Aqueveque et al. 2018; Carrera-Játiva et al. 2023) may be because this rodent is more exposed to various helminths due to its generalist habitat and diet, as well as its high densities (Spotorno et al. 2000). A prevalence greater than 20 % has been reported in few Chilean rodent species (Landaeta-Aqueveque et al. 2018; Grandón-Ojeda et al. 2022).

In general, high prevalence heterogeneity (I2) values were found in the different parasites. This finding may be due to multiple factors, such as differences between the sampled locations and in the diagnostic methods used. Regarding the latter, 2 meta-analyses that produced higher I2 values (Leptospira spp. and protozoa) use at least 3 different diagnostic methods. It is also worth mentioning that the measure used to estimate heterogeneity has some limitations because high I2 values (< 90) have been reported as frequent in prevalence meta-analyses (Migliavaca et al. 2022).

Based on the results of this study, we propose the following: 1) increase the study of viruses with zoonotic potential, since most studies have focused on Orthohantavirus using serological techniques only; 2) advance the use of molecular techniques for more accurate identification of bacteria, protozoa, and helminths; 3) increase the study of ticks in the olive grass mouse and deepen the potential role of this species and its mites in relation to rickettsial infections such as scrub typhus; 4) from an ecological perspective, it is necessary to carry out comparative studies of the parasite-olive grass mouth association in relation to changes in land use, habitat fragmentation, and other human disturbances in environments. In addition, eco-epidemiological studies should include the rodent community coexisting with the olive grass mouse. This will generate additional knowledge about the pathogens circulating at the community level and deepen our understanding of differences in prevalence between rodent species. The above will provide additional information that can be used in meta-analysis studies and more robust spatiotemporal comparisons of the olive grass mouse and its parasites, thereby helping to better understand the epidemiological role of this rodent related to various zoonotic pathogens.

Acknowledgements

H. Henríquez thanks the Agencia Nacional de Investigación y Desarrollo (ANID), Chile for the National Ph. D. Grant (Folio 21221386). The authors wish to thank two anonymous reviewers for their comments and suggestions, which significantly improved the manuscript. M. E. Sánchez-Salazar translated the manuscript into English.

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Associated editor: Itandehui Hernández Aguilar.

Submitted: February 15, 2024; Reviewed: May 29, 2024.

Accepted: June 7, 2024; Published on line: June 18, 2024.

Appendix 1

References included in the meta-analyses, but not found in the article.

Espinoza‐Rojas, H., et al. 2021. Survey of Trichinella in American minks (Neovison vison Schreber, 1777) and wild rodents (Muridae and Cricetidae) in Chile. Zoonoses and Public Health 68:842-848.

Fernández, J., et al. 2008. Identificación de Hantavirus Andes en Rattus norvegicus. Archivos de Medicina Veterinaria 40:295-298.

Galuppo, S., et al. 2009. Predominance of Trypanosoma cruzi genotypes in two reservoirs infected by sylvatic Triatoma infestans of an endemic area of Chile. Acta Tropica 111:90-93.

Landaeta-Aqueveque, C., et al. 2014. First record of Litomosoides pardinasi (Nematoda: Onchocercidae) in native and exotic rodents from Chile. Revista Mexicana de Biodiversidad 85:1032-1037.

Murúa, R., et al. 2003. Síndrome pulmonar por Hantavirus: situación de los roedores reservorios y la población humana en la Décima Región, Chile. Revista Médica de Chile 131:169-176.

Ortiz, J., et al. 2004. Hantavirus en roedores de la Octava Región de Chile. Revista Chilena de Historia Natural 77:151-156.

Rozas, M., et al. 2007. Coexistence of Trypanosoma cruzi genotypes in wild and periodomestic mammals in Chile. American Journal of Tropical Medicine and Hygiene 77:647.

Sánchez, R., et al. 2020. Rodents as potential reservoirs for Borrelia spp. in northern Chile. Revista Brasileira de Parasitologia Veterinaria 29:e000120.

Torres-Pérez, F., et al. 2004. Peridomestic small mammals associated with confirmed cases of human hantavirus disease in southcentral Chile. The American Journal of Tropical Medicine and Hygiene 70:305-309.

Appendix 2

PRISMA flowchart showing the item selection steps

Appendix 3

Forest plot of the prevalence of ANDV by host subgroup (Abrothrix olivacea, Oligoryzomys longicaudatus, and other rodents). Each green square represents prevalence. Black lines represent the 95 % confidence intervals. Gray diamonds represent the mean prevalence of each subgroup.

Appendix 6

Forest plot of the prevalence of ectoparasites by host subgroup (Abrothrix olivacea and other rodents). Each green square represents prevalence. Black lines represent the 95 % confidence intervals. Gray diamonds represent the mean prevalence of each subgroup.

Table 1. Micro- and macroparasites reported in Abrothrix olivacea in Chile. *Considered zoonotic; **belongs to a genus that includes zoonotic species; ~~ vectors "cf." indicates a comparison between the individual in question and the referred taxon, based on the consistency of observable characters, although these characters are not sufficiently numerous. 1Virus tentatively named 'Olivaceaous mouse morbilivirus'. 2Toxoplasmatinae gen. sp. P99 is genetically distinct at the genus level within the subfamily, so it is believed to be a new genus. Regions: AP (Arica and Parinacota), TA (Tarapacá). AN (Antofagasta), AT (Atacama), CO (Coquimbo), VA (Valparaíso), ME (Metropolitan), OH (O'Higgins), MA (Maule), ÑU (Ñuble), BB (Biobío), AR (Araucanía), LR (Los Ríos), LL (Los Lagos), AY (Aysén), MG (Magallanes). Diagnostic method: 1 (Strip Immunoblot Assay), 2 (ELISA), 3 (RNAseq), 4 (PCR), 5 (Immunohistochemistry), 6 (Dark-Field Microscopy), 7 (Bacterial culture), 8 (Microscopy agglutination test), 9 (Argentica stain), 10 (Indirect immunofluorescence), 11 (Immunoperoxidase), 12 (Telemann stool test for parasites), 13 (Ziehl-neelsen stool test for parasites), 14 (Flotation stool test for parasites), 15 (Light microscopy), 16 (Scanning Microscopy), 17 (Microscopy), 18 (Mini-FLOTAC stool test for parasites), 19 (Isolation by necropsy).

	Parasite	Location (regions)	Diagnostic method	Reference
Viruses	ANDV*	CO, VA, ME, OH, MA, ÑU, BB, AR, LR, LL, AY	1, 2	Rubio et al. 2019; Torres-Pérez et al. 2019; Medina et al. 2009; Toro et al. 1998
	R.O. Morbilivirus1	LR	3	Debat 2022
Bacteria
	Mycoplasma sp.**	LR	4	Alabí et al. 2020.
	Leptospira spp. (L. borgpetersenii and L. interrogans)*	LR	4-5, 8-9	Luna et al. 2020; Riedemann and Zamora 1982
	Leptospira spp.*	ME, LR	4, 6-7	Correa et al. 2017; Zamora et al. 1999
	Leptospira interrogans*	ME, LR	8, 10-11	Perret et al. 2005; Zamora et al. 1995
	Yersinia enterocolitica*	LR	7	Zamora et al. 1998
	Bartonella spp.**	ME, OH, AR	4	Sepúlveda-García et al. 2023
Protozoans
Metamonad	Giardia sp.**	MA	12	Infante et al. 2022
Apicomplexa	Cryptosporidium sp.**	MA	13	Infante et al. 2022
	Coccidia	LL	14	Carrera-Játiva et al. 2023
	Hepatozoon spp.	AP, TA, AT, CO, LR, LL	4	Alabí et al. 2021; Merino et al. 2009; Santodomingo et al. 2022
	Sarcocystis sp.	OH, ÑU, AR, LR, LL	4	Oyarzún-Ruiz et al. 2023
	Toxoplasmatinae2	OH, ÑU, AR, LR, LL	4	Oyarzún-Ruiz et al. 2023
Euglenozoa	Trypanosoma cruzi*	CO, VA, ME	4	Botto-Mahan et al. 2010; Botto-Mahan et al. 2020; Correa et al. 2015; Ihle-Soto et al. 2019; Oda et al. 2014; Rozas et al. 2007
Amoebozoa	Amoeba sp.	LL	14	Carrera-Játiva et al. 2023
Ectoparasites
Acari	Ixodes sp.~~	ME	15	Muñoz-Leal et al. 2019
	Ixodes abrocomae	CO	4-16	Guglielmone et al. 2010
	Herpetacarus eloisae~~	LL, AY	17	Silva-de la Fuente et al. 2021; Silva-de la Fuente et al. 2023
	Herpetacarus sp.~~	LL	17	Acosta-Jamett et al. 2020
	Paratrombicula goffi	LL, AY	17	Silva-de la Fuente et al. 2021; Silva-de la Fuente et al. 2023
	Paratrombicula neuquenensis~~	LL, AY	17	Silva-de la Fuente et al. 2023
	Paratrombicula sp.~~	LL	17	Acosta-Jamett et al. 2020
	Quadraseta chiloensis	LL, AY	17	Silva-de la Fuente et al. 2021; Silva-de la Fuente et al. 2023
	Quadraseta sp.	LL	17	Acosta-Jamett et al. 2020
	Argentinacarus expansus~~	LL, AY	17	Silva-de la Fuente et al. 2023
	Ornithonyssus sp.~~	MA	17	Veloso-Frías et al. 2019
	Androlaelaps sp.	MA	17	Veloso-Frías et al. 2019
Siphonaptera	Ctenoparia inopinata~	TA, AN, AT, CO, VA, ME, OH, MA, ÑU, BB, AY, MG	17	Alarcón 2003; Bazán-León et al. 2013; Moreno-Salas et al. 2020
	Ctenoparia jordani	TA, AT, CO, VA, ME, OH, MA, ÑU, AY, MG	17	Moreno-Salas et al. 2020
	Ctenoparia topalli	TA, AT, CO, VA, ME, OH, MA, ÑU, AY, MG	17	Moreno-Salas et al. 2020
	Ectinorus angularis	AN, AT, CO, ME, BB	17	Bazán-León et al. 2013
	Ectinorus cocyti	TA, AT, CO, VA, ME, OH, MA, ÑU, AY, MG	17	Moreno-Salas et al. 2020
	Hectopsylla cypha	AN, AT, CO, ME, BB	17	Bazán-León et al., 2013
	Hectopsylla spp.~~	TA, AT, CO, VA, ME, OH, MA, ÑU, AY, MG	17	Moreno-Salas et al. 2020
	Neotyphloceras chilensis~~	TA, AN, AT, CO, VA, ME, OH, MA, ÑU, BB, AY, MG	17	Bazán-León et al. 2013; Moreno-Salas et al. 2020
	Neotyphloceras crassispina~~	TA, AN, AT, CO, VA, ME, OH, MA, ÑU, BB, AY, MG	17	Alarcón 2003; Bazán-León et al. 2013; Moreno-Salas et al. 2020
	Neotyphloceras pardinasi~~	TA, AT, CO, VA, ME, OH, MA, ÑU, AY, MG	17	Moreno-Salas et al., 2020
	Sphinctopsylla ares~~	TA, AN, AT, CO, VA, ME, OH, MA, ÑU, BB, AY, MG	17	Alarcón 2003; Bazán-León et al. 2013; Moreno-Salas et al. 2020
	Tetrapsyllus corfidii	TA, AN, AT, CO, VA, ME, OH, MA, ÑU, BB, AY, MG	17	Bazán-León et al. 2013; Moreno-Salas et al. 2020
	Tetrapsyllus rhombus~~	TA, AN, AT, CO, VA, ME, OH, MA, ÑU, BB, AY, MG	17	Alarcón 2003; Bazán-León et al. 2013; Moreno-Salas et al. 2020
	Tetrapsyllus tantillus~~	TA, AN, AT, CO, VA, ME, OH, MA, ÑU, BB, AY, MG	17	Alarcón 2003; Bazán-León et al. 2013; Moreno-Salas et al. 2020
	Tetrapsyllus amplus	TA, AT, CO, VA, ME, OH, MA, ÑU, AY, MG	17	Moreno-Salas et al. 2020
	Nosopsyllus fasciatus~~	TA, AT, CO, VA, ME, OH, MA, ÑU, BB, AY, MG	17	Alarcón 2003; Moreno-Salas et al. 2020
	Plocopsylla wolffsohni	BB	17	Alarcón 2003
	Chiliopsylla allophyla~~	BB	17	Alarcón 2003
	Agastopsylla boxi	TA, AT, CO, VA, ME, OH, MA, ÑU, AY, MG	17	Moreno-Salas et al. 2020
	Listronius spp.	TA, AT, CO, VA, ME, OH, MA, ÑU, AY, MG	17	Moreno-Salas et al. 2020
	Leptopsylla segnis~~	TA, AT, CO, VA, ME, OH, MA, ÑU, AY, MG	17	Moreno-Salas et al. 2020
Phthiraptera	Hoplopleura andina	VA	17	González-Acuña et al. 2003; González-Acuña et al. 2005
Hemiptera	Mepraia paraprática~~	TA	4	Quiroga et al. 2022
Endoparasites
Acanthocephala	Moniliformis sp.**	MA, LL	12, 18	Carrera-Játiva et al. 2023; Riquelme et al. 2021
Nematode	Physaloptera sp.	MA, LL	12, 19	Carrera-Játiva et al. 2023; Riquelme et al. 2021
	Physaloptera calnuensis	CO, VA, ME	19	Landaeta-Aqueveque et al. 2007; Landaeta-Aqueveque et al. 2018
	Syphacia sp.**	MA, LL	12, 18, 19	Carrera-Játiva et al. 2023; Riquelme et al. 2021
	Syphacia obvelata*	CO, VA, ME	19	Landaeta-Aqueveque et al. 2007; Landaeta-Aqueveque et al. 2018
	Syphacia phyllotios	CO	19	Yáñez-Meza et al. 2019
	Capillaria sp.**	MA, ME	12, 19	Landaeta-Aqueveque et al. 2007; Riquelme et al. 2021
	Capillariidae**	LL	18	Carrera-Játiva et al. 2023
	Helminthoxys gigantea	CO	19	Yáñez-Meza et al. 2019
	Trichuris sp.**	LL	18, 19	Carrera-Játiva et al. 2023
	cf. Anatrichosoma sp.**	CO, VA, ME	19	Landaeta-Aqueveque et al. 2018
	Strongyloides sp.**	LL	18	Carrera-Játiva et al. 2023
	Pterygodermatites sap.	CO, VA, ME	19	Landaeta-Aqueveque et al. 2007; Landaeta-Aqueveque et al. 2018
	Protospirura sp.	LL	19	Carrera-Játiva et al. 2023
	Heterakis spumosa	CO, VA, ME	19	Landaeta-Aqueveque et al. 2007; Landaeta-Aqueveque et al. 2018
	Gongylonema sp.**	CO	19	Yáñez-Meza et al. 2019
Cestode	Hymenolepis sp.**	CO, VA, ME, MA	12, 19	Landaeta-Aqueveque et al. 2007; Landaeta-Aqueveque et al. 2018; Riquelme et al. 2021
	Rodentolepis sp.**	LL	18	Carrera-Játiva et al. 2023

Table 1. Continuation...

Figure 1. Distribution map of the number of studies by region in Chile.

Table 2. Prevalence, confidence intervals, and heterogeneity index of micro- and macroparasites in Abrothrix olivacea and other rodents in Chile. Prevalence and confidence intervals are expressed in %. *This rodent was only separated in the hantavirus analysis because it is recognized as the main reservoir of the virus.

	Abrothrix olivacea	Other rodents	Oligoryzomys longicaudatus*	Total
Virus (ANDV)	0.4 (0.0-1.5), I2 = 84 %	0.9 (0.4-1.5), I2 = 24 %	4.6 (3.2-6.2), I2 = 44 %	1.6 (0.6-3.0), I2 = 92 %
Bacteria	32.3 (9.5-59.3), I2 = 95 %	22.3 (12.2-34.3), I2 = 95 %	--	25.3 (15.2-36.7), I2 = 95 %
Leptospira	37.8 (12.6-66.9), I2 = 96 %	30.0 (13.6-49.5), I2 = 97 %	--	33.8 (20.4-48.6), I2 = 96 %
Protozoans	25.7 (12.6-41.1), I2 = 97 %	22.9 (12.4-35.3), I2 = 98 %	--	24.1 (15.7-33.4), I2 = 97 %
Trypanosoma cruzi	46.6 (31.9-61.5), I2 = 76 %	40.7 (29.6-52.3), I2 = 95 %	--	43.0 (34.1-52.1), I2 = 92 %
Ectoparasites	10.8 (5.2-17.8), I2 = 95 %	14.6 (8.0-22.7), I2 = 95 %	--	12.6 (8.2-17.8), I2 = 95 %
Helminths	8.4 (5.2-12.2), I2 = 94 %	2.8 (1.1-4.9), I2 = 91 %	--	5.4 (3.6-7.4), I2 = 93 %
Coproparasitic helminths	4.5 (2.6-7.0), I2 = 89 %	1.3 (0.3-2.8), I2 = 76 %	--	2.9 (1.7-4.4), I2 = 87 %
Helminths necropsy	14.6 (6.2-25.5), I2 = 95 %	5.2 (1.7-10.1), I2 = 93 %	--	9.7 (5.5-14.8), I2 = 95 %

Table 3. Meta-regression of prevalence according to parasite group. Abrothrix olivacea is considered a reference category. c analysis using stool tests for parasites; n analysis through isolates from specimen necropsy.

ANDV	Parameter	Standard error	Z value	P value
Intercept	0.09	0.02	4.64	< 0.0001
Oligoryzomys longicaudatus	0.17	0.03	6.26	< 0.0001
Other rodents	0.05	0.03	1.81	0.070
Bacteria
Intercept	0.63	0.10	6.21	< 0.0001
Other rodents	-0.13	0.14	-0.96	0.336
Leptospira
Intercept	0.68	0.12	5.54	< 0.0001
Other rodents	-0.09	0.17	-0.51	0.611
Protozoans
Intercept	0.55	0.08	7.02	< 0.0001
Other rodents	-0.05	0.11	-0.44	0.659
Protozoa
Intercept	0.57	0.10	5.95	< 0.0001
Other rodents	-0.04	0.13	-0.33	0.742
Trypanosoma cruzi
Intercept	0.75	0.08	9.49	< 0.0001
Other rodents	-0.06	0.11	-0.56	0.578
Ectoparasites
Intercept	0.35	0.05	6.65	< 0.0001
Other rodents	0.05	0.07	0.73	0.465
Helminths
Intercept	0.31	0.03	9.44	< 0.0001
Other rodents	-0.10	0.05	-2.17	0.030
Helminths (c)
Intercept	0.22	0.03	8.39	< 0.0001
Other rodents	-0.08	0.04	-1.98	0.048
Helminths (n)
Intercept	0.40	0.05	7.96	< 0.0001
Other rodents	-0.13	0.07	-1.84	0.066

Appendix 4

Forest plot of the prevalence of bacteria by host subgroup (Abrothrix olivacea and other rodents). Each green square represents prevalence. Black lines represent the 95 % confidence intervals. Gray diamonds represent the mean prevalence of each subgroup.

Appendix 5

Forest plot of the prevalence of protozoa by host subgroup (Abrothrix olivacea and other rodents). Each green square represents prevalence. Black lines represent the 95 % confidence intervals. Gray diamonds represent the mean prevalence of each subgroup.

Appendix 7

Forest plot of the prevalence of helminths by host subgroup (Abrothrix olivacea and other rodents). Each green square represents prevalence. Black lines represent the 95 % confidence intervals. Gray diamonds represent the mean prevalence of each subgroup.