THERYA, 2025, Vol. 16(2):283-292
Gastrointestinal parasitism in nine-banded armadillos
(Dasypus novemcinctus) under different hunting pressures
in the Reserva Natural del Bosque Mbaracayú, Paraguay.
Martin Kowalewski1*, M. Belén Natalini1, Cathia Coronel2,3, Sol Gennuso1, Myriam C. Velázquez3, Federico Pontón1,
Berta Martínez4, Fredy Ramírez Pinto3, M. Cecilia Ezquiaga5, Sixto Fernandez3, and Thomas R. Gillespie6
1Estación Biológica Corrientes (EBCo), CECOAL-CONICET (Consejo Nacional de Investigaciones Científicas y Técnicas), Corrientes, Argentina. Email: martinkow@gmail.com (M-K); belennatalini@gmail.com (MB-N); solgennuso@gmail.com (S-G); federicoponton@gmail.com (FC-P).
2Centro para el Desarrollo de Investigación Científica, Asunción, Paraguay. Email: cathiacoronel@gmail.com (C-C).
3Fundación Moisés Bertoni, Asunción, Paraguay. Email: mvelazquez@mbertoni.org.py (MC-V); framirezpy@gmail.com (F-RP); sxtfernandez@gmail.com (S-F).
4Universidad Nacional de Canindeyú, Paraguay. Email: bertamartinezc18@gmail.com (B-M).
5Centro de Estudios Parasitológicos y de Vectores (CEPAVE) - CONICET, UNLP, La Plata, Argentina. Email: cecilia@cepave.edu.ar (MC-E).
6Departments of Environmental Sciences and Environmental Health, Emory University and Rollins School of Public Health, Atlanta, Georgia, USA Email: thomas.gillespie@emory.edu (TR-G)
*Corresponding author: https://orcid.org/0000-0002-6737-3771
The aim of this study was to characterize the gastrointestinal parasitic fauna of nine-banded armadillos (Dasypus novemcinctus) of the Reserva Natural del Bosque Mbaracayú (RNBM), Canindeyú, Paraguay, and to evaluate how human activities, particularly hunting, may influence parasitism patterns and zoonotic risk. We recovered 15 distinct parasites from fresh fecal samples of 73 individuals of D. novemcinctus including Protozoa: Eimeria sp., Giardia sp., coccidia (Family Adeleidae), unidentified coccidia; Cestodes: Hymenolepis sp.; Nematodes: Trichuris sp., Trichostrongyloidea, Cruzia sp., Ancylostoma sp., Uncinaria sp., Aspidodera sp., Strongyloides sp. and Acanthocephala. We report high prevalence of potentially zoonotic parasites including Ancylostoma, Trichuris, Uncinaria and Coccidia. We also found an increase in parasite richness in areas where hunting was reported vs. areas without hunting and the same trend for Nematode probability of presence, especially during the cold dry season. These results suggest that human activities may be affecting patterns of parasitism in wild armadillos. In addition, traditional consumption of these animals in this region may present a route of human exposure to zoonotic parasites.
El objetivo de este estudio fue caracterizar la fauna parasitaria gastrointestinal de armadillos de nueve bandas (Dasypus novemcinctus) de la Reserva Natural del Bosque Mbaracayú (RNBM), Canindeyú, Paraguay y evaluar cómo las actividades humanas, particularmente la caza, pueden influir en los patrones de parasitismo y el riesgo de zoonosis. Recuperamos 15 parásitos distintos de muestras fecales frescas de 73 individuos de D. novemcinctus, incluidos protozoos: Eimeria sp., Giardia sp., coccidias (familia Adeleidae), coccidias no identificadas; Cestodos: Hymenolepis sp.; Nematodos: Trichuris sp., Trichostrongyloidea, Cruzia sp., Ancylostoma sp., Uncinaria sp., Aspidodera sp., Strongyloides sp. y Acanthocephala. Informamos una alta prevalencia de parásitos potencialmente zoonóticos, incluidos Ancylostoma, Trichuris, Uncinaria y Coccidia. También encontramos un aumento en la riqueza de parásitos en áreas donde se reportó caza versus áreas sin caza y la misma tendencia en la probabilidad de presencia de nematodos, especialmente durante la estación fría y seca. Estos resultados sugieren que las actividades humanas pueden estar afectando los patrones de parasitismo en los armadillos salvajes. Además, el consumo tradicional de estos animales en esta región puede presentar una vía de exposición humana a parásitos zoonóticos.
Keywords: Zoonoses, helminths, prevalence, human-wildlife interface
© 2025 Asociación Mexicana de Mastozoología, www.mastozoologiamexicana.org
Introduction
Nine-banded armadillos (Dasypus novemcinctus Linnaeus, 1758) extend from Colombia and Venezuela to Argentina (MacDonald 1985; Redford and Eisenberg 1992; Loughry et al. 2015; Barthe et al. 2024). In Paraguay, they occur throughout the country, but are considerably rare in arid areas (Smith 2007; De la Sancha et al. 2017; Smith and Ríos 2018). They are nocturnal, generalist insectivores that spend their days in self-dug burrows, often shared by multiple individuals from the same family group (Taber 1945; Nowak 1991; Parera 2002; Smith 2007). In Paraguay, many human cultural groups use this species as a food resource including Aché, Mbyá, and Guaraní indigenous groups and farmers from rural settlements (McBee and Baker 1982; Esquivel 2001; Neris et al. 2002; Hill et al. 2003; Smith 2007; Morales 2009).
Although the parasites of D. novemcinctus are well characterized at the northern extent of their range (i.e.: Chandler 1946; Radomski et al. 1991; Varela-Stokes et al. 2008; Jimenez-Ruiz et al. 2013), previous research in the southern extent of their range is more limited. For example, Fujita et al. (1995) reported some nematodes recovered from the gastrointestinal tract of just two specimens (Dept. of San Pedro, Central Paraguay): Ancylostoma sp., Aspidodera esperanzae (possibly A. binansata -Jimenez-Ruiz et al. 2006), Moeniggia complexus, M. pintoi and an unidentified species of Heterakinae. Navone (1990) recorded the nematode Aspidodera vazi (Aspidoderidae) and the cestode Mathevotaenia surinamensis (Anoplocephalidae) in the Paranaense and Chaco regions of Argentina, with Aspidodera fasciata (Aspidoderidae) additionally recorded in the Chaco region. In Santa Cruz, Bolivia Jimenez-Ruiz et al. (2003) reported Aspidoderid nematodes Aspidodera binansata (n=173) and Lauroia bolivari sp. nov (n=22) from the gastrointestinal tract of one individual. Hoppe and do Nascimento (2007) reported at least 13 species of nematodes recovered from the gastrointestinal tract of nine individuals in Pantanal (Mato Grosso, Brazil). Also, in Bolivia, Deem et al. (2009) reported the following parasites from 22 individuals: Heterakidae, Aspidodera sp., Trichuris sp., Adelaidae (protozoan), and Strongylus sp.
The species can coexist with humans in rural areas because of its largely nocturnal habits and large litter size but has declined over much of its range in eastern Paraguay because of destruction of optimal habitats and hunting pressure (Smith 2007). Armadillos of the studied species may present zoonotic pathogens thus hunting and human consumption of these species or sharing edge habitats may have a significant public health impact (Stallknecht et al. 1987; Yaeger et al. 1988; Truman 2005; Deem et al. 2009).
The aim of the current study was twofold: 1) to begin the characterization of the gastrointestinal parasitic fauna of nine-banded armadillos (Dasypus novemcinctus) of the Reserva Natural del Bosque Mbaracayú (RNBM), Canindeyú, Paraguay; and 2) to explore differences in parasite diversity and prevalence between areas exposed to human activities (i.e.: traditional and illegal hunting) and more remote, undisturbed areas within the reserve. In addition to determining the parasites present and their prevalence in Dasypus novemcinctus within the RNBM, we also aimed to determine if the parasite communities and/or the prevalence of specific taxa differed in armadillos from hunted areas compared to those from non-hunted areas. By comparing infection patterns across zones with contrasting hunting pressure, we also aimed to evaluate the influence of these human practices on parasite in armadillos, as well as the potential zoonotic risks for individuals involved in their hunting and consumption (Medkour et al. 2020; Inada 2023; Mozer and Prost 2023; Salvarani et al. 2025).
Methods
Study site. Our study was conducted as part of a larger research program to investigate infectious diseases dynamics in the protected area Mbaracayú Forest Nature Reserve (RNBM) and its surroundings in Paraguay. Specifically, this study was done at the Reserva Natural del Bosque Mbaracayú (RNBM), (24° 00’ - 24° 15’ S, 55° 00’ - 55° 32’ W), Depto. Canindeyú, Paraguay. The RNBM represents a c. 640 km2 protected area composed mostly of Upper Parana Atlantic Forest. Mbaracayú was established in 1991 and is privately managed by a Paraguayan non-governmental organization (Fundación Moises Bertoni). The study area is also within a UNESCO biosphere reserve, which includes the protected area Mbaracayú Forest Nature Reserve (RNBM), as a core preserved area.
The climate is warm, humid, and temperate, with an average annual temperature of 21–22 °C and annual rainfall ranging from 1,650 to 1,800 mm, peaking between October and March. Based on the classification by Owen et al. (2010), the year is divided into three seasons: transitional (T: February–May), dry (D: June–September), and wet (W: October–January). Frosts can occasionally occur from June to October. The landscape consists of a mosaic of natural forest and savanna ecosystems, including High Forest, Medium and Low Forests with bamboo, wetlands, cerrado, and native grasslands. Forests range from mature stands with open understories to dense, bamboo-dominated areas (Chusquea ramosissima, Guadua spp.), while savannas include Cerrado formations and grassy riparian zones, contributing to the area’s ecological complexity (FMB/BM 2005).
Sampling was conducted in two types of areas within the RNBM: (1) areas under direct human influence, such as zones where hunting occurs, and (2) remote, minimally disturbed forested areas where no hunting occurs (Figure 1). One of the human-impacted sites was located near Arroyo Bandera, the closest community to the reserve. In this region, indigenous communities have legal authorization to practice traditional hunting, while illegal hunting by non-Indigenous individuals has also been reported both along the edges and within the interior of the reserve. These designations were based on reports from rangers responsible for protection and monitoring of the RNBM. In the hunting zones, the most predominant domestic species observed was the dog (Canis lupus familiaris), which is commonly used by hunters as a companion and a hunting aid during their excursions into the forest (Kowalewski et al. 2019).
Study subjects. Sample collection occurred from June 2012 to August 2013, and from September 2016 to June 2017. For subsequent analyses, samples were classified according to the three seasonal periods previously defined (transitional, dry and wet). Fresh fecal samples were collected from D. novemcinctus, primarily from individuals captured by hand by Aché technicians with expertise in traditional hunting practices. Additional samples were obtained opportunistically from fresh feces found in the field, attributed to non-captured individuals. Once captured, each armadillo was placed in an individual breathable bag and transported to our nearby base camp where our veterinarian performed a health assessment and obtained a fecal sample from each individual. All armadillos were weighed while in the bag and then immobilized using ketamine-zolazepam for physical examination and collection of biologic samples. After sampling, armadillos were released at the site of capture. All fecal samples were preserved in 10% buffered formalin in 20 ml plastic tubes and labeled. We did not repeat capture sites, and armadillos were marked previous release to avoid recapture.
Parasitological Analyses. Fecal samples were examined using direct microscopic examination. Techniques to recover eggs included Willis, Sheather and zinc sulphate methods (MAAF 1986; Thienpont et al. 1986; Foreyt 1989) at the Parasitological Laboratory of the Centro para el Desarrollo de la Investigación Científica (CEDIC, Asunción, Paraguay). Slides (18 mm x 18 mm) prepared by each method were observed under a light microscope (LEICA DM500) after the addition of a drop of Lugol’s iodine and photographed with a Leica camera (ICC50W). Parasites were identified based on egg, larvae, or cyst coloration, shape, contents, and size following Anderson et al. (1974), Khalil et al. (1994), Lainson and Shaw (1982), Mehlhorn et al. (1992), and Navone (1986; 1987; 1988, 1990). Each slide was scanned thoroughly under both 10x and 40x objective lenses to confirm presence or absence of helminth eggs and larvae and protozoan cysts. Representatives of each parasite species recovered per sample were measured at 400x to the nearest 0.1 µm with the Leica Application Suite software (2016) and representative parasite species were photographed. Morphological examination of eggs and oocysts allowed diagnosis to the genus level or higher levels for some parasites. Thus, the taxonomic level achieved through morphological identification of parasite eggs or oocysts depended on the parasite, as some genera had distinctive eggs enabling genus-level diagnosis, while others could only be identified at higher taxonomic levels, such as the superfamily Trichostrongyloidea
Statistical analyses. We estimated the prevalence (as the proportion of individuals positive for a given parasite divided by the total number of armadillos examined) and richness (as the number of the parasite species/group present per individual). Using Generalized Linear Models (GLM), we analyzed the effects of environmental, hunting and sex of the individual on richness and parasite occurrence (presence/absence of parasites) (Table 1). For these analyses we divided data into broad categories, presence of nematodes, presence of cestodes and presence of protozoan parasites (Model 2.a; 2.b; 2.c respectively). Full models were then restricted by removing non-significant terms in a stepwise backward elimination manner, using likelihood ratio tests, until we obtained a final model that contained only significant variables. GLM were fit by maximum likelihood (Laplace Approximation), using the function “glm” from the MASS package (Venables and Ripley 2002). Poisson (log) distribution was used for the richness model and Binomial (logit) distribution for the prevalence set of models. For the first model we verified Poisson distribution of dependent variable “richness” graphically (plot distribution of cumulative frequencies and qq-plot). The assumptions of normally distributed and homogeneous residuals (Pearson residuals) were checked by visually inspecting qq-plots and the residuals plotted against the fitted values. We used R version 3.2.1 (The R Project for Statistical computing, www.r-project.org) for the analyses, and statistical significance was set at a p-value of 0.05. for all interpretations.
Compliance with ethical standards. We have followed ARRIVE guidelines and all procedures complied with relevant legislation, including the U.K. Animals (Scientific Procedures) Act (1986), EU Directive 2010/63/EU, and the NIH Guide for the Care and Use of Laboratory Animals. We used the minimum number of animals required for statistically valid results. The study adhered to Paraguayan law (N° 96/92 on wildlife), with permits for scientific collection (N°06/2012 and 270/2016) granted by Ministry of the Environment and Sustainable Development (MADES-Paraguay in Spanish).
Results
We collected a total of 78 fresh fecal samples. Of these, 66 correspond to D. novemcinctus captured by hand and the other 12 samples correspond to fresh fecal samples of non-captured armadillos. Of the 66 weighed armadillos, 32 were females and 34 males. There were no differences in weight between females (4.6 ±0.8 kg) and males (4 ± 1.2 kg) (p>0.05). All captured individuals showed good health status based on physical examination.
We recovered 15 distinct parasite taxa from 73 (out of 78) individuals including Protozoa: Eimeria sp., Giardia sp., coccidia (Family Adeleidae), unidentified coccidia; Cestodes: Hymenolepis sp.; Nematodes: Trichuris sp., Trichostrongyloidea, Cruzia sp., Ancylostoma sp., Uncinaria sp. Aspidodera sp., Strongyloides sp. and Acanthocephala (possible Travassosia sp.) (Table 2, Figure 2). The total prevalence of gastrointestinal parasites was 93.6%. Both Ancylostoma sp. and Trichuris sp. presented the highest prevalence: 67.1% and 90.4% respectively. When coccidia parasites were pooled, the general prevalence reached 71.2% (Table 2). The number of distinct parasites per individual ranged from 1-9.
Parasite richness was higher in sites with hunting and during the dry season compared to sites without hunting and the wet season, respectively (Figure 3). Neither sex nor precipitation were significant contributors to the final model (Table 3). Presence / absence models showed a trend toward hunting pressure being associated with a higher probability of the presence of nematodes (Table 4). Null models were selected in protozoan and cestode models.
Discussion
In this study we recovered 15 distinct parasites from 73 wild D. novemcinctus in Reserva Natural del Bosque Mbaracayú (RNBM) including taxa known to be prevalent in species of the Order Xenarthra, including coccidia of the Families Adeleidae and Eimeriidae (Lange and Wittenstein 2001; Ezquiaga et al. 2014), nematodes of the genera Aspidodera (Vicente 1966; Vicente et al. 1997; Hoppe and Nascimento 2007) and Cruzia (Fujita et al. 1995; Vicente et al. 1997; Hoppe and Nascimento 2007), and an acanthocephala, presumed to be Oligacanthorhynchus (syn, Travassosia) sp., (Travassos 1917; Lent and Freitas 1938; Smales 2007; Gomez-Puerta 2012). Notably, Giardia spp., Hymenolepis spp., and Uncinaria spp. were recorded for the first time in this host species, expanding the known parasitic diversity of D. novemcinctus.
The richness found in our research is similar to that recorded by Hoppe and do Nascimento (2007) in Brazil, and higher than that recorded by Fujita et al. (1995) in Paraguay, Navone (1990) in Argentina and Deem et al. (2009) in Bolivia. However, methodological differences and ecological contexts across these studies must be considered. For instance, Fujita et al. (1995) examined only two museum specimens, with no specific environmental information available. Navone (1990) analyzed 17 gastrointestinal tracts from individuals collected in temperate grasslands and subtropical regions of Buenos Aires and Corrientes provinces. Hoppe and do Nascimento (2007) also relied on gastrointestinal tract analysis, although detailed ecological context was not provided. In contrast, Deem et al. (2009) analyzed fecal samples from 22 individuals collected within the Gran Chaco region of Bolivia, a dry, thorny forest and savanna ecosystem. These variations among samples (feces vs. gastrointestinal tracts), sample size, and habitat likely influence parasite detection and diversity, and should be carefully considered when comparing across studies. Nevertheless, a summary of parasite records across different countries is presented in Table 5, to contextualize our findings and highlight general patterns and knowledge gaps.
Overall parasite richness was higher in areas with hunting, with the strongest effect occurring during the dry season (from June to September). We found a trend toward the same pattern when comparing the probability of parasite presence / absence (i.e., higher probability of presence in areas with hunting and during the dry season) for nematodes, but not cestodes or protozoa. These results suggest that human activities within the reserve may alter patterns of parasitism in wild armadillos. One possible explanation for our results is that in the areas where illegal hunting is known to occur, poachers establish camps and use their dogs for hunting (Ramirez Pinto, pers. comm.). Both, dogs and humans, defecate on the ground and could contribute to a high rate of soil and grass contamination with infective parasitic stages. Studies in nearby communities have reported a high diversity of gastrointestinal parasites in domestic dogs, including Uncinaria stenocephala, Strongyloides stercoralis, Ancylostoma caninum, Toxocara canis, Cystoisospora sp., Trichuris vulpis, Giardia duodenalis, and Taenia sp. (Martínez 2019). Notably, some of these parasites—such as Giardia, Trichuris, and Uncinaria—were also detected in armadillos from hunting areas, suggesting potential overlap. Given their feeding habits, armadillos may be especially exposed to fecal-oral transmission routes and could act as accidental or paratenic hosts for these parasites. Furthermore, the higher parasite richness observed during the colder winter months (June to August) in comparison with the rest of the year, may be related to lower temperatures and food availability, which could compromise immune function and increase host susceptibility to infection. Additionally, some nematodes, such as Trichuris, produce highly resistant eggs that can remain viable in the environment under these conditions (Traversa et al. 2011). These survival traits, combined with seasonal nutritional stress and immune compromise, may contribute to sustained environmental infective pressure and help explain the observed increase in parasite richness during the dry season. Concerning cestodes and protozoa, there was not a significant association or trend between these groups of parasites and the independent variables. In general, cestodes have indirect cycles and use insects as intermediate hosts. Although we did not collect specific data on the diet of the captured armadillos, field observations confirm the presence of termite mounds in several areas where individuals were captured. As insectivorous mammals, armadillos are known to dig into the soil and termite nests in search of food, which could provide them access not only to termites and ants but also to insect larvae developing underground. In the case of protozoa is probably that the presence of some parasites (e.g., Adeleidae) is associates to the mechanical passage of oocysts from infected prey through the gastrointestinal tract of the nine - banded armadillos, rather than reflecting true parasitism (Lange and Wittenstein 2001; Ezquiaga et al. 2009).
Although we recovered some taxa with zoonotic potential, such as nematodes of the genera Ancylostoma and Trichuris (at high prevalence), Uncinaria (at moderate prevalence), and protozoa such as Giardia to a lesser extent (Table 2; Figure 2), it is important to note that the presence of these genera does not necessarily imply a direct zoonotic risk. Most species within these groups can infect both animals and humans; however, not all species possess the ability to infect humans or cause disease. Future molecular analyses are essential to identify the specific species involved and to accurately assess their zoonotic potential.
Understanding these risks is particularly important given that the traditionally consumption of these animals by legal and illegal hunters may represent a potential route of human exposure to zoonotic parasites- either through ingestion of contaminated meat or through exposure to body fluids and feces during hunting and preparation. Moreover, Ache hunters eat part of the game during their hunting expeditions (Ramirez Pinto, pers. comm.), which increases the risk of transmission when meat is consumed raw or undercook, or when handling and transportation practices do not follow food safety practices (Thompson 2013; Van Vliet et al. 2017).
Based on these practices, we suggest that exposure to parasites may occur not only through ingestion of infected tissue, but also via handling of raw meat or cross-contamination during food preparation. The risk may arise from indirect contact with fecal material present in the environment during traditional hunting practices. In such contexts, hunters—often accompanied by dogs—frequently dig into burrows to manually extract the animals, increasing the likelihood of exposure to soil and fecal contamination (Natalini pers. comm.) This, combined with the consumption practices described above, creates multiple potential pathways for zoonotic transmission.
Our study represents a descriptive approximation, and we do not aim to explore transmission dynamics in depth. While we acknowledge that environmental conditions such as soil moisture can influence the transmission of hookworms and other parasites, our current data are insufficient to make conclusive statements in this regard. Nonetheless, this is indeed an important aspect that warrants further investigation through targeted ecological and longitudinal studies.
Considering these findings, which highlight the possibility of zoonotic transmission, future studies employing molecular analyses and opportunistic necropsies are needed to improve our classification of the gastrointestinal parasites of armadillos. Mechanisms to reduce zoonotic risks identified may promote human health and local support for conservation.
Conclusions
We characterized the gastrointestinal parasitic fauna of nine-banded armadillos (Dasypus novemcinctus) recovering 15 distinct parasites from fresh fecal samples of 73 individuals at the Reserva Natural del Bosque Mbaracayú (RNBM), Canindeyú, Paraguay. We also report an increase in parasite richness in areas where hunting was reported in contrast to areas without hunting and the same trend for Nematode probability of presence. These findings suggest that human activities may be affecting patterns of parasitism in wild armadillos.
Acknowledgments
We want to give special thanks to the Ache people that were involved in this project: Pastor Mbepurangi, Ambrosio Jaguagi, Felipe Bejarogi, Tito Jaguagi, Germino Nuñez. MK, MBN and MCE are members of CONICET-Argentina. MK thanks BK for helping him to understand solitary terrestrial displacements. The authors acknowledge Laura Rodríguez of the Moisés Bertoni Foundation for designing and preparing the map. This work was supported by the Consejo Nacional de Ciencia y Tecnología (CONACYT)-Programa PROCIT and co-financed by CONACYT - Programa PROCIENCIA (Paraguay) with the support of FEEI, through the following projects: “INV 36 Estudio y prevención de zoonosis en la Reserva de Biosfera del Bosque Mbaracayú, Paraguay” y “14INV052 La salud de los ecosistemas, transmisión de enfermedades y alteración del hábitat en la Reserva de Biosfera del Bosque Mbaracayú”. MK and TG are thankful to Emory University, the Fulbright Program from the United States and CONICET from Argentina. The content of this publication is the exclusive responsibility of the authors, and under no circumstances it should be considered as reflecting the opinion of CONACYT.
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Associated editor: Alina Gabriela Monroy Gamboa
Submitted: October 1, 2024; Reviewed: November 11, 2024;
Accepted: May 20, 2025; Published on line: May 30, 2025
DOI:10.12933/therya-25-6160 ISSN 2007-3364
Figure 1. Location of Dasypus novemcinctus sampling sites with and without hunting within the Reserva Natural del Bosque Mbaracayú, Paraguay.
Table 1. Variables summary for GLM model.
|
Model |
Variable |
Definition |
|
Dependent variables |
||
|
1 |
Richness |
Count (min= 1; max=9) |
|
2. a |
Presence nematodes |
Binary response |
|
2. b |
Presence cestodes |
Binary response |
|
2. c |
Presence protozoa |
Binary response |
|
Independent Variables |
||
|
Hunting |
Categorical variable, |
|
|
Sex |
Categorical variable, |
|
|
Environmental |
*Season |
Categorical variable, 3 levels |
|
Monthly total precipitation |
Continuous variable |
Table 2. Prevalence of parasites recovered in fresh fecal samples from 73 individual Dasypus novemcinctus (out of 78 individuals examined) at Reserva Natural del Bosque de Mbaracayú, Paraguay. Twenty individuals are from hunting areas and 53 individuals are from areas without hunting.
|
Parasite/group |
Prevalence (%) |
||
|
Protozoa |
Total |
Hunting |
Without Hunting |
|
Giardia sp.* |
6.9 |
21.4 |
3.4 |
|
Eimeria sp. (Coccidia) |
5.5 |
14.3 |
3.4 |
|
Unidentified Coccidia |
49.3 |
71.4 |
44.1 |
|
Adeleidae (Coccidia) |
16.4 |
42.9 |
10.2 |
|
Cestoda |
|||
|
Unknown Cestode 1 |
5.5 |
14.3 |
3.4 |
|
Hymenolepis sp.* |
4.1 |
7.2 |
3.4 |
|
Nematoda |
|||
|
Strongyloides sp. |
6.9 |
7.2 |
6.8 |
|
8.2 |
21.4 |
5.1 |
|
|
Trichuris sp. |
90.4 |
100 |
88.1 |
|
Uncinaria sp.* |
20.6 |
42.9 |
15.3 |
|
Ancylostoma sp. |
67.1 |
57.1 |
69.5 |
|
Cruzia sp. |
4.1 |
7.1 |
3.4 |
|
Aspidodera sp. |
15.0 |
21.4 |
13.6 |
|
Unknown Nematode 1 |
1.4 |
7.1 |
0 |
|
Acanthocephala (possibly Oligacanthorhynchus) |
1.4 |
7.1 |
0 |
(*) indicate new records for the species.
Figure 2. Parasites recovered from fecal samples of Dasypus novemcinctus within the Reserva Natural del Bosque Mbaracayú, Paraguay. (a) Ancylostoma sp. 1 (b) Uncinaria sp. (c) Strongyloides sp. (d) Trichostrongyloidea (e) Cruzia sp. (f) Trichuris sp. (g) Hymenolepis sp. (h) Aspidodera sp. (i) Coccidio, Adeleidae Family (j) Acanthocephala (possible Travassosia sp.) (k) Cestode 1 Scale bar is 20 μm.
Richness
N = Sites without hunting; Y = Sites with hunting.
Figure 3. Richness GLM results. Blue line represents expected richness and gray boxes are standard deviations according to GLM. Seasons are T, transitional (February–May); D, dry (June–September); W, wet (October–January) following Owen et al. (2010).
Table 3. GLM results, independent variables estimate of the model selected for richness.
|
Model |
Estimate |
Std. Error |
z value |
Pr(>|z|) |
|
(intercept) |
1.16 |
0.12 |
9.27 |
0.001*** |
|
hunting (Yes) |
0.59 |
0.15 |
3.81 |
0.001*** |
|
season (T) |
-0.37 |
0.16 |
-2.33 |
0.01* |
|
season (W) |
-0.37 |
0.16 |
-2.32 |
0.02* |
Table 4. GLM results, independent variables estimate of the model selected for presence / absence of nematode.
|
Model |
Estimate |
Std. Error |
z value |
Pr(>|z|) |
|
(intercept) |
1.7 |
0.54 |
3.14 |
0.01** |
|
hunting (Yes) |
18.8 |
2614.2 |
0.007 |
0.99 |
Season
Table 5. Reported parasites of Dasypus novemcinctus across its distribution range.
|
Parasite |
Sample |
Location |
Reference |
|
Brachylaemus virginianus, Oncicola canis, Hamanniella sp., Aspidodera fasciata, Physocephalus sp., Ascarops sp. |
GI tracts (N= 8) |
USA |
|
|
Aspidodera vazi (Aspidoderidae) and the cestode Mathevotaenia surinamensis (Anoplocephalidae), Aspidodera fasciata (Aspidoderidae) |
GI tracts (N=17) |
Argentina |
|
|
Macracanthorhynchus ingens, Centrorhynchus sp., Physaloptera sp. |
GI tract (N=117) |
USA |
|
|
Ancylostoma sp., Aspidodera esperanzae (possibly A. binansata -Jimenez-Ruiz et al. 2006), Moeniggia complexus, M. pintoi, unidentified species of Heterakinae. |
GI tract (N= 2) |
Paraguay |
|
|
Aspidodera binansata, Lauroia bolivari sp. nov |
GI tract (N=1) |
Bolivia |
|
|
Aspidodera sogandaresi, Aspidodera vazi, Aspidodera binansata, Lauroia trinidadensis, L. trinidadensis, |
GI tract (N= 13) |
Mexico Panama USA |
|
|
Macielia macieli, M. flagellata, Moennigia moennigi, M. complexus, M. pintoi, M. littlei, Delicata variabilis, Hadrostrongylus speciosum, Strongyloides ratti, Aspidodera fasciata, A. binansata, A. vazi, Cruzia spp. |
GI tract (N= 9) |
Brazil |
|
|
Aspidodera cf. soganderesi, Macracanthorhynchus ingens |
GI tract (N=?) |
USA |
|
|
Heterakidae, Aspidodera sp., Trichuris sp., Adelaidae, Strongylus sp. eggs |
Fecal samples (N= 22) |
Bolivia |
|
|
Mathevotaenia sp., Centrorhynchus sp. |
GI tract (N= 30) |
Brazil |
|
|
Cestoda, Trematode eggs, Trichostrongylidae and Ascarididae eggs, Capillaria sp. |
Fecal samples (N= 4) |
Brazil |
GI= gastrointestinal, N= number of samples collected.