image/svg+xmlEste artículo es publicado por la revista Neotropical Helminthology de la Facultad de Ciencias Naturales y Matemática, Universidad Nacional Federico Villarreal, Lima, Perú auspiciado por la Asociación Peruana de Helmintología e Invertebrados Afines (APHIA). Este es un artículo de acceso abierto, distribuido bajo los términos de la licencia Creative Commons Atribución 4.0 Internacional (CC BY 4.0) [https:// creativecommons.org/licenses/by/4.0/deed.es] que permite el uso, distribución y reproducción en cualquier medio, siempre que la obra original sea debidamente citada de su fuente original.ISSN Versión impresa 2218-6425ISSN Versión Electrónica 1995-1043Neotropical Helminthology, 2022, 16(2), jul-dic:161-172.ORIGINAL ARTICLE / ARTÍCULO ORIGINAL1Laboratório de Biologia e Parasitologia de Mamíferos Silvestres Reservatórios, Instituto Oswaldo Cruz, Fundação Oswaldo Cruz, Av. Brasil 4365, 21040-360, Rio de Janeiro, RJ, Brasil.2 Campus Fiocruz Mata Atlântica, Av. Sampaio Corrêa s/n, Fundação Oswaldo Cruz, 22713-560, Rio de Janeiro, RJ, Brasil.3Programa de Pós-Doutorado Júnior, Conselho Nacional de Desenvolvimento Científico e Tecnológico, Brasília, Brasil.4 Programa de Pós-Doutorado Nota 10 – 2021, Fundação Carlos Chagas Filho de Amparo à Pesquisa do Estado do Rio de Janeiro, Brasil.5 Programa de Pós-Graduação em Biodiversidade e Saúde, Instituto Oswaldo Cruz, Fundação Oswaldo Cruz, Av. Brasil, 4365, 21040-360, Rio de Janeiro, RJ, Brasil.6Programa INOVA IOC, Vice-presidência de Pesquisa e Coleções Biológicas, Fundação Oswaldo Cruz, Av. Brasil 4365, 21040-360, Rio de Janeiro, RJ, Brasil7Departamento de Parasitologia Animal, Universidade Federal Rural do Rio de Janeiro, km 07, BR-465, 23890- 000, Seropédica, RJ, Brasil.*Corresponding author: rgentile@ioc.fiocruz.brRosana Gentile: https://orcid.org/0000-0001-9160-1204Sócrates Fraga da Costa Neto: https://orcid.org/0000-0003-2813-525XThiago dos Santos Cardoso: https://orcid.org/0000-0003-2724-6038Raquel Gonzalez Boullosa: https://orcid.org/0000-0002-7560-4277Carla Elizabeth Macabu: https://orcid.org/0000-0003-2109-0933Raquel de Oliveira Simões: https://orcid.org/0000-0001-5130-3341Arnaldo Maldonado Júnior: https://orcid.org/0000-0003-4067-86601*2,31,41,5,6Rosana Gentile; Sócrates Fraga da Costa-Neto; Thiago dos Santos Cardoso; Raquel Gonzalez Boullosa; 17 1Carla Elizabeth Macabu; Raquel de Oliveira Simões& Arnaldo Maldonado-JúniorNeotropical Helminthology161DOI: http://dx.doi.org/10.24039/rnh20221621451HELMINTHS OF SMALL MAMMALS IN AN ATLANTIC FOREST BIOLOGICAL STATION IN RIO DE JANEIRO, BRAZILHELMINTOS DE PEQUEÑOS MAMÍFEROS EN UNA ESTACIÓN BIOLÓGICA DEL BOSQUE ATLÁNTICO EN RÍO DE JANEIRO, BRASILHELMINTOS DE PEQUENOS MAMÍFEROS EM UMA ESTAÇÃO BIOLÓGICA DA MATA ATLÂNTICA NO RIO DE JANEIRO, BRASILDDDDDDABSTRACTInterface areas between urban and sylvatic environments increase the contact between humans and wild animals, and may favour the transmission of zoonoses. The aim of this study was to describe the helminth fauna of a small mammal community in an urban-sylvatic interface area of the Brazilian Atlantic Forest. art. 1=10-24art. 2=26-41Art 3 =42-51art. 4=52-59art. 5=60-65art 6=66-79art. 7=80-91art 8 =92-100art 9=102-111nota 1=112-116nota 2=118-123 rev =124-134
image/svg+xmlTwenty helminth species were recovered in six species of small mammals. Parasite sharing was observed in two helminth species among the marsupials. This study is the first report of a helminth infection for the marsupialMonodelphis americana(Müller, 1776). This is the first report of the nematodes Aspidodera raillieti Travassos, 1913, Viannaia hamataTravassos, 1914 and Trichuris sp. parasitizing the marsupial Marmosa paraguayana (Tate, 1931). None of the helminth species found has been reported to infect 162Neotropical Helminthology, 2022, 16(2), jul-dicKeywords: Marsupials – Prevalence – Rodents – Synanthropic animals – Urbanization – Wild animalsThe continuous process of urbanization and expansion of human activities in forested areas disrupts natural habitats and may change the patterns of species diversity and abundance. In disturbed ecosystems, generalist species, such as the common opossum Didelphis aurita (Wied-Neuwied, 1826), or opportunistic species, such as some sigmodontine rodents, are favoured in relation to specialist species (Gentile et al., 2018). They may have their densities increased and may INTRODUCTIONbecome abundant in rural and peridomicile areas (D'Andrea et al., 2007; Kajin et al., 2008). The occurrence of synanthropic species, such as the common rat Rattus norvegicusBerkenhout, 1769, and the black rat Rattus rattus(Linnaeus, 1758), is also observed in these areas. Such changes in the demography and distribution of natural populations, especially in the areas of the interface between urban and rural or urban and forest environments, directly influence the transmission dynamics of parasites (Kruse et al., 2004). These interface areas may also harbour invasive species. The presence of invasive species and their parasites Gentile et al.Las áreas de interfaz entre ambientes urbanos y selváticos aumentan el contacto entre humanos y animales salvajes y pueden favorecer la transmisión de zoonosis. El objetivo de este estudio fue describir la helmintofauna de una comunidad de pequeños mamíferos en un área de interfaz urbano-selvática de la Mata Atlántica brasileña. Se recuperaron veinte especies de helmintos en seis especies de pequeños mamíferos. Se observó el intercambio de parásitos en dos especies de helmintos entre los marsupiales. Este estudio es el primer reporte de una infección por helmintos para el marsupial Monodelphis americana(Müller, 1776). Este es el primer reporte de los nematodos Aspidodera raillietiTravassos, 1913, Viannaia hamataTravassos, 1914 y Trichurissp. parasitando al marsupial Marmosa paraguayana(Tate, 1931). No se ha informado que ninguna de las especies de helmintos encontradas infecte a los humanos.RESUMENPalabras clave: Animales sinantrópicos – Animales salvajes – Marsupiales – Prevalencia – Roedores – Urbanización Áreas de interface entre ambientes urbanos e silvestres aumentam o contato entre os seres humanos e os animais silvestres, podendo favorecer a transmissão de zoonoses. O objetivo deste estudo foi descrever a helmintofauna de uma comunidade de pequenos mamíferos em uma área de interface urbano-silvestre da Mata Atlântica brasileira. Vinte espécies de helmintos foram recuperadas em seis espécies de pequenos mamíferos. Compartilhamento de parasitos foi observado em duas espécies de helmintos entre os marsupiais. Este estudo é o primeiro relato de infecção helmíntica pelo marsupial Monodelphis americana(Müller, 1776). Este é o primeiro relato dos nematoides Aspidodera raillieti Travassos, 1913, Viannaia hamataTravassos, 1914 e Trichurissp. parasitando o marsupial Marmosa paraguayana (Tate, 1931). Nenhuma das espécies de helmintos encontrada já foi relatada infectando seres humanos.RESUMOPalavras-chave: animais sinantrópicos – Animais silvestres – marsupiais – prevalência – roedores – urbanizaçãoart. 1=10-24art. 2=26-41Art 3 =42-51art. 4=52-59art. 5=60-65art 6=66-79art. 7=80-91art 8 =92-100art 9=102-111nota 1=112-116nota 2=118-123 rev =124-134
image/svg+xmlmay affect the local communities and the host-parasite relationships (Lucio et al., 2021).The occurrence of human infection by helminth parasites from wild animals has been attributed to the increase of interface areas between urban or rural areas with forest fragments or reserves, where species of reservoir animals occur. Knowledge of the helminth species that parasitize wild mammals in these areas is essential for understanding their role as an etiological and/or zoonotic agent and, consequently, the potential risk to public and animal health (Bezerra-Santos et al., 2020), due to the continuous process of urbanization and human advance on forest areas.There are several studies of helminth faunas of small mammals in the Brazilian Atlantic Forest (Gomes et al., 2003; Maldonado et al., 2006; Cardoso et al., 2016; Boullosa et al., 2020; Lucio et al., 2021). However, there is a large gap concerning helminth studies in forested reserves close to urban centres (Costa-Neto et al., 2019; Boullosa et al., 2021). The present study is complementary to Costa-Neto et al. (2019), which studied the structure of the helminth metacommunity of the common opossum Didelphis auritain three localities, including the locality of the present study. The aim of the present study was to describe the helminth fauna of a small mammal community and the helminth population parameters in an urban-sylvatic interface area of the Atlantic Forest in Brazil. Study areaThis study is part of a comprehensive project on biodiversity and environmental health aiming to understand the role of mammals in the transmission cycles of parasites and zoonotic agents at the FIOCRUZ Atlantic Forest Biological Station (EFMA) (Estação Biológica FIOCRUZ Mata Atlântica) and its surroundings, including areas of Pedra Branca State Park (PEPB) (Parque Estadual da Pedra Branca), in the city of Rio de Janeiro, southeast Brazil. EFMA is located on the eastern slope of the Pedra Branca massif in the western zone of the city of Rio de Janeiro, which is the second most populous city in Brazil. This area 163MATERIAL AND METHODSwas previously occupied by sugarcane mills and coffee farms, with the construction of residences and roads. In 2003, the area was incorporated into the Fundação Oswaldo Cruz (FIOCRUZ). From then on, the master plan of the institution delimited nonedification areas and established a more effective process of environmental protection and recovery aiming to mitigate the effects of anthropic impacts. The area is formed by a mosaic of landscapes on the border of the forest reserve, including sites with vegetation in an advanced stage of ecological succession, transition sites with intermediate and initial stages of reforestation, and sites with consolidated urbanization.Pedra Branca State Park, adjacent to EFMA, is the largest urban forest in the Americas and one of the largest remnants of the Atlantic Forest biome in the state of Rio de Janeiro. It covers an area of 12,492 ha, and the predominant phytophysiognomy is ombrophilous dense Atlantic Forest vegetation. The region's climate is humid mesothermic, with hot, rainy summers and mild winters.The samplings were made in different types of habitat representing preserved forest areas (22°56'45"S, 43°25'00"W), transition areas between urban and forested environments, including reforestation areas (22°56'29"S, 43°24'25”W), and peridomicile areas (22°56'18”S, 43°24'11”W). See Gentile et al. (2018) for a more detailed description of the study area. The areas of preserved forest were located distant from human dwellings and, for the most part, above 100 m elevation. These areas have canopy heights ranging from 10 to 30 m and irregular slopes, ranging from flat to steep. The transition areas were located between the peridomicile areas and the preserved forest, with a canopy height ranging from 6 to 20 m and a flat slope. The peridomicile areas were located in the backyards of the houses, with a lower canopy ranging from 6 to 10 m, predominance of shrubs and small trees, presence of flooded areas and slope varying from flat to moderate.Small mammal samplingSmall mammal captures were carried out from July 2012 to April 2014 every four months and in July, October and November 2017 for five consecutive nights in the three types of habitat. Mammals were collected using Tomahawk® (16 × 5 × 5 inches) and Sherman® (3 × 3.75 × 12 inches) live-traps. Neotropical Helminthology, 2022, 16(2), jul-dicHelminths of small mammals in Rio de Janeiroart. 1=10-24art. 2=26-41Art 3 =42-51art. 4=52-59art. 5=60-65art 6=66-79art. 7=80-91art 8 =92-100art 9=102-111nota 1=112-116nota 2=118-123 rev =124-134
image/svg+xml164Two transects of twenty points were established in each habitat. On each transect, two traps were placed on the ground at each point. All traps were baited with a mixture of peanut butter, banana, oatmeal and bacon. The total trapping effort was 9,120 trap-nights. The animals were anesthetized and submitted to euthanasia for helminth search. The mammals were identified by external morphology, cranial morphology and cytogenetic analyses, when necessary. The mammalian specimens were submitted to taxidermy and deposited in the mammalian collection of the National Museum of the Federal University of Rio de Janeiro voucher: 83156 - 83163; 83167 - 83172.Collection and identification of helminthsViscera, thoracic and abdominal cavities, and musculature were examined for helminths. The organs were separated in Petri dishes with saline solution (0.85% NaCl) and dissected using a stereoscopic microscope. The nematodes were fixed in AFA solution (93 parts 70% ethanol, 5 parts 0.4% formol, and 2 parts 100% acetic acid) and heated to 65°C. Some specimens were stored in 70% ethanol for further molecular analysis. Trematodes were fixed in the same solution under compression, and the cestodes and acanthocephalans were kept in distilled water for relaxation of the musculature (Amatoet al., 1991). Nematodes were diaphanized with lactophenol or glycerinated alcohol. Trematodes, cestodes and acanthocephalans were stained with Langeron's carmine or Delafield's hematoxylin, differentiated with 0.5% hydrochloric acid, dehydrated in a crescent alcoholic series, diaphanized in methyl salicylate and mounted in Canada balsam for permanent preparation (Amato et al., 1991). The specimens were analysed under a light microscope (Axio Scope A1 light microscope – Zeiss, Göttingen, Germany) coupled to an Axio Cam MRc digital camera.The specific morphological aspects used to identify the specimens were according to Andersonet al.(2009), Gibbons (2010), Gibson et al. (2002), Jones et al.(2005), Khalil et al.(1994), Travassos (1937), Vicente et al.(1997) and Yamaguti (1961). The specimens were deposited in the scientific collection of helminths at the Laboratory of Biology and Parasitology of Wild Mammal Reservoirs - IOC/FIOCRUZ-RJ using the same deposit numbers of their respective hosts.Helminth population parametersPopulation parameters were calculated for each helminth species in each host species according to Bush et al.(1997). The average abundance was considered as the total number of helminths divided by the number of host individuals analysed. Mean intensity was calculated as the total number of helminths divided by the number of animals infected. Prevalence was considered to be the ratio between the number of infected animals and the total number of animals analysed. These three parameters were calculated for each helminth species considering each host species. Mean helminth species richness was calculated as the average of the helminth species richness found in each infracommunity (community harboured by a single host) for each host species.A bipartite network plot to illustrate host-parasite interactions between species (Poulin, 2010) was carried out between the small mammals and the helminth species. This plot was built based on a matrix of the presence/absence of each helminth species in each host species using the bipartite package (Dormann et al., 2008).Ethic aspectsAnimals were captured under authorization of the Brazilian Government's Chico Mendes Institute for Biodiversity and Conservation (ICMBIO, licence number 13373) and the Environmental Institute of Rio de Janeiro State (INEA, licence number 020/2011). All procedures followed the guidelines for the capture, handling, and care of animals of the Ethical Committee on Animal Use of the Oswaldo Cruz Foundation (CEUA, licence numbers LW81/12, and LW-39/14). Biosafety procedures and personal safety equipment were used during all procedures involving animal handling and biological sampling.Seven species of small mammals were captured, four of which were marsupials, Didelphis aurita (Wied Neuwied, 1826) (45), Metachirus myosurus (Temminck, 1924) (1), Marmosa paraguayanaTate, 1931 (5) and Monodelphis americana (Müller, 1776) (9) (Didelphimorphia, Didelphidae), and three were rodents, Akodon RESULTSNeotropical Helminthology, 2022, 16(2), jul-dicGentile et al.art. 1=10-24art. 2=26-41Art 3 =42-51art. 4=52-59art. 5=60-65art 6=66-79art. 7=80-91art 8 =92-100art 9=102-111nota 1=112-116nota 2=118-123 rev =124-134
image/svg+xml165Table 1.Mean abundance, intensity (± SD), prevalence (95% CI) and site of infection of the helminths of marsupials at FIOCRUZ Atlantic Forest Biological Reserve, Rio de Janeiro, Brazil. (-) indicates absence of the parasite species.Neotropical Helminthology, 2022, 16(2), jul-dicHelminths of small mammals in Rio de Janeiroart. 1=10-24art. 2=26-41Art 3 =42-51art. 4=52-59art. 5=60-65art 6=66-79art. 7=80-91art 8 =92-100art 9=102-111nota 1=112-116nota 2=118-123 rev =124-134
image/svg+xmlTable 2.Mean abundance, intensity (± SD), prevalence (95% CI) and site of infection of the helminths of rodents at FIOCRUZ Atlantic Forest Biological Reserve, Rio de Janeiro, Brazil. (-) indicates absence of the parasite species.166Neotropical Helminthology, 2022, 16(2), jul-dicGentile et al.art. 1=10-24art. 2=26-41Art 3 =42-51art. 4=52-59art. 5=60-65art 6=66-79art. 7=80-91art 8 =92-100art 9=102-111nota 1=112-116nota 2=118-123 rev =124-134
image/svg+xml167cursor (Winge, 1887) (7), Oligoryzomys nigripes(Olfers, 1818) (9) (Rodentia, Sigmodontinae), and the sinantropic rodent Rattus rattus(Linnaeus, 1758) (7) (Rodentia, Muridae). Twenty species of helminths were collected from the small mammal community. Thirteen species were collected from D. aurita, three from M. paraguayana, one from M. americana, two from A. cursor, two from O. nigripesand two from R. rattus(Table 1). No helminth was found in the marsupial M. myosurus.Forty-five individuals of D. auritawere analysed, and all were infected by at least one helminth species. The mean species richness was 5.4 and varied from 1 to 9 among the infracommunities. The highest abundances and intensities were observed for the nematodes Viannaia hamata Travassos, 1914 (Nematoda: Viannaiidae Durette-Desset & Chabaud, 1981), followed by Cruzia tentaculata(Rudolphi, 1819) Travassos (1917) (Nematoda: Kathlaniidae Travassos, 1918) and Travassostrongylus orloffi Travassos, 1935 (Nematoda: Viannaiidae Durette-Desset & Chabaud, 1981) (Table 1). The highest prevalence was recorded for C. tentaculata, T. orloffiand Turgida turgida (Rudolphi, 1819) Travassos, 1919 (Nematoda: Physalopteridae Leiper, 1908) (Table 1). M. paraguayanahad two specimens infected among the five analysed. The helminth species found were Aspidodera raillieti Travassos, 1913 (Nematoda: Aspidoderidae (Freitas, 1956), Trichurissp. 1, and V. hamata, the latter showing the highest prevalence and abundance in this host (Table 1). The mean species richness was 1. Only two specimens among nine M. americanawere infected by a single helminth species, V. hamata(Table 1).Considering rodents, seven individuals of A. cursorwere analysed, and two of them were infected by helminths, the cestode Rodentolepis akodontisRêgo, 1967 (Platyhelminthes, Hymenolepididae) and the nematode Trichurissp. 2 (Table 2). All helminths presented low abundances and intensities. The mean species richness was 0.43. Four individuals among nine O. nigripesanalysed were infected. The helminth species found were Guerrerostrongylus zetta(Travassos, 1937), Sutton & Durette-Desset, 1991 (Nematoda, Heligmonellidae) and Stilestrongylus lanfrediaeSouza et al., 2008 (Nematoda, Heligmonellidae), both with low abundance and intensity (Table 2). The mean species richness was 0.55. Seven individuals of the synanthropic rodent R. rattus were analysed, and two were infected. Two helminth species were found: Nippostrongylus brasiliensis(Travassos, 1914) (Nematoda, Heligmonellidae) in a single host and Syphacia muris(Yamaguti, 1935) (Nematoda, Oxyuridae) in two hosts, the latter with the largest abundance and intensity found among rodent hosts (Table 2). The mean species richness was 0.43.The bipartite network plot between small mammals and helminths showed little parasite sharing among host species (Fig. 1). The nematode V. hamatawas shared between the marsupials D. aurita, M. paraguayanaand M. americana(Fig. 1). D. auritaand M. paraguayanashared the nematode A. raillieti(Figure 1). No other helminth species sharing was observed (Figure 1).The marsupial D. auritahad the highest helminth species richness when compared to the other host species. This fact can be due to the generalist habit of this species, to its vagility (Cáceres & Monteiro-Filho, 2001), which may have contributed to a greater acquisition of parasites in different environments, to its larger body size in relation to the other mammals and to the larger number of specimens captured and analysed. The low helminth species richness found for all the other host species may be due to the high disturbance of the local environment, resulting in helminth species whose eggs may be more resistant in the soil or species less sensitive to environmental alterations. Other studies found that habitat fragmentation and disturbances could negatively affect the life cycles of some species of parasites, limiting the occurrence of some species '(Lafferty & Kuris, 1999; Cardoso et al., 2016). However, we cannot discard the low sample sizes of these mammals.This study is the first report of a helminth infection for the marsupialM. americana, which was infected by V. hamata.Furthermore, this is the first report of the nematodes A. raillieti, V. hamataand Trichuris sp. parasitizing M. paraguayana, DISCUSSIONNeotropical Helminthology, 2022, 16(2), jul-dicHelminths of small mammals in Rio de Janeiroart. 1=10-24art. 2=26-41Art 3 =42-51art. 4=52-59art. 5=60-65art 6=66-79art. 7=80-91art 8 =92-100art 9=102-111nota 1=112-116nota 2=118-123 rev =124-134
image/svg+xml168increasing its helminth fauna to six species and adding this mammal as a new host for these helminths. Little is known about the helminth fauna of M. paraguayana(Santos-Rondon et al., 2012). In Brazil, only two nematode species have been reported in this host, the oxyurid Gracilioxyuris agilisisFeijó, Torres, Maldonado Jr & Lanfredi, 2008 in the Cerrado (Neotropical savannah), state of São Paulo (Santos-Rondon et al., 2012), and the filariid Litomosoides barrettiMuller, 1980 in the state of Bahia (Muller, 1980). The cestode Mathevotaenia bivittata(Janicki, 1904) has also been reported in this host in Argentina (Campbell et al., 2003). The helminths of D. auritaare discussed in detail in Costa-Neto et al. (2019).Regarding the sigmodontine rodent A. cursor, which occurs only in the Brazilian Atlantic Forest up to 1,170 m high (Geise, 2012), the cestode R. akodontishad been previously reported in this host Simões et al.(2011), as well as a nematode of the genus Trichuris(T. navonae) by Lucio et al. (2021). The helminth species richness reported in the present study was very low compared to previous reports of the helminth fauna of this rodent in the Figure 1. A bipartite network plot indicating the small mammal – helminth associations at FIOCRUZ Atlantic Forest Biological Reserve, Rio de Janeiro, Brazil. Neotropical Helminthology, 2022, 16(2), jul-dicGentile et al.art. 1=10-24art. 2=26-41Art 3 =42-51art. 4=52-59art. 5=60-65art 6=66-79art. 7=80-91art 8 =92-100art 9=102-111nota 1=112-116nota 2=118-123 rev =124-134
image/svg+xml169Atlantic Forest. Simões et al. (2011) found nine species in forest fragments, and Lucio et al. (2021) found eight species in open matrix areas, both in the state of Rio de Janeiro.The two nematodes found in O. nigripesare commonly reported in other studies of the helminth fauna of this sigmodontine rodent (Pintoet al., 1982; Gomeset al., 2003; Simões et al., 2011; Werk et al., 2016; Panisse et al., 2017; Boullosa et al., 2020; Cardoso et al., 2020). Likewise, the two nematodes recovered from the synanthropic rodent R. rattushave also been reported for this host in other studies (Gómez-Muñoz et al., 2018; Panti-May et al., 2020). In Argentina, Muñoz et al. (2018) found higher values of abundance and prevalence of N. brasiliensisin R. rattusthan in this study. Panti-May et al. (2019) found low values of intensity, abundance and prevalence for this helminth in a region of Mexico. These authors also found high prevalence of S. murispositively associated with the abundance of R. rattus.The helminth species were shared only within a single taxonomic group (Tribe Didelphini), with no record of parasite sharing between rodents and marsupials. The two helminths (V. hamataand A. raillieti) shared by the marsupial species can be considered core species in the present study, with high prevalence and abundances.Indeed, A. raillietiand V. hamatapresented prevalence greater than 50% in D. aurita. This indicates that the three didelphids, D. aurita, M. paraguayanaand M. americana, which share the environment, also share some helminth species. Jiménezet al. (2011), studying helminth component communities of didelphid marsupials from several localities, suggested that sympatric species of marsupials shared more species of parasites than parasite communities occurring in conspecific marsupials from different localities. Our results corroborate that study with regard to the local scale.The results indicated little sharing of parasite species in these small mammal-helminth associations. Although these mammals have been reported as reservoirs of zoonotic pathogens in the study area(Berbigier et al., 2021), and some of them may harbour helminths with zoonotic potential (Bezerra-Santos et al., 2020; Meerburg et al., 2009; Araújo et al., 2014), none of the identified helminth species are known to infect humans.We would like to thank the staff and students of Laboratório de Biologia e Parasitologia de Mamíferos Silvestres Reservatórios, Laboratório de Hantaviroses e Rickettsioses, Laboratório de Biologia de Tripanossomatíedeos and Laboratório de Pesquisa Clínica em Dermatozoonose from FIOCRUZ who helped in the field work; the coordinator of FIOCRUZ Atlantic Forest Campus, Gilson Antunes, for providing local facilities and supporting the general project; the equip of FIOCRUZ Atlantic Forest Campus, especially to Ricardo Moratelli, Marta L. Brandão and José L.P. Cordeiro for local facilities and help in the field work; to Marcelo F. Freitas for helping in the field work in all campaigns; to Cibele R. Bonvicino for rodent identification, Fabiana P. Caramaschi for marsupial identification and Daniela Dias for the curator of the collected animals. This project was financially supported by Conselho Nacional de Desenvolvimento Científico e Tecnológico-CNPq (AMJ, grant number 306352/2014-1), (RG, researcher fellowship number 304355/2018-6),Fundação Carlos Chagas Filho de Amparo à Pesquisa do Estado do Rio de Janeiro-FAPERJ (RG grant number E-26/111.296/2014 and E-26/010.001597/2019), and Instituto Oswaldo Cruz (FIOCRUZ), Fundação Oswaldo Cruz (RGB, PhD scholarship), (TSC postdoctoral fellowship).ACKNOWLEDGEMENTSAmato, JFR, Boeger, W & Amato, SBB, 1991. Protocolos para laboratório: coleta e processamento de parasitos de pescado. Imprensa Universitária, UFRRJ, Rio de Janeiro.Anderson, RC, Chabaud, AG & Willmott, S. 2009. Keys to the nematode parasites of vertebrates. CABI International, Wallingford.Araújo EO, Mendes, MM, Langone, PQ & Müller, BIBLIOGRAPHIC REFERENCESNeotropical Helminthology, 2022, 16(2), jul-dicHelminths of small mammals in Rio de Janeiroart. 1=10-24art. 2=26-41Art 3 =42-51art. 4=52-59art. 5=60-65art 6=66-79art. 7=80-91art 8 =92-100art 9=102-111nota 1=112-116nota 2=118-123 rev =124-134
image/svg+xml170G. 2014. The helminth parasites of Rattus rattus(Linnaeus, 1758) of urban, intermediate and rural environments in southern brazil. Neotropical Helminthology, vol. 8, pp. 19-22. Berbigier, AP, Barros, JHS, Pontes, ES, Lisboa, CV., Gentile, R, Xavier, SCC, Jansen, AM & Roque, ALR. 2021. Trypanosomatid richness in wild and synanthropic small mammals from a biological station in Rio de Janeiro, Brazil.Pathogens, vol. 10, 1442.Bezerra-Santos MA, Fontes CS, Nogueira BCF, Yamatogi RS, Ramos RAN, Galhardo JA, Furtado LFV, Rabelo EML, Araújo JV & Campos AK. 2020. Gastrointestinal parasites in the opossum Didelphis aurita: are they a potential threat to human health?Journal of Parasitic Diseases, vol. 44, pp. 355–363. Boullosa, RG, Cardoso, TS, Costa-Neto, SF, Teixeira, BR, Freitas, TPT, Maldonado-Júnior, A & Gentile, R. 2020. Helminth community structure of three sigmodontine rodents in the Atlantic Forest, southern Brazil. Oecologia Australis, vol. 24, pp. 577–589. Boullosa, RG, Costa-Neto, SF, Morgado, L, Maldonado Jr, A & Gentile, R. 2021. A longitudinal survey of gastrointestinal parasites of the black-eared opossum Didelphis auritaat an urban-sylvatic interface area in Southeast Brazil: a morphological and ecological characterization based on helminth eggs. Parasitology Research, vol.120, pp.3815–3825.Bush, AO, Lafferty, KD, Lotz, JM & Shostak, AW. 1997. Parasitology meets ecology on its own terms: Margolis et al. revisited. Journal of Parasitology, vol. 83, pp. 575–583.Cáceres, NC & Monteiro-Filho, ELA. 2001. Food habits, home range and activity of Didelphis aurita(Mammalia, Marsupialia) in a forest fragment of Southern Brazil. Studies on Neotropical Fauna & Environment, vol. 36, pp. 85–92.Campbell, ML, Gardner, SL, & Navone, GT. 2003. A new species of Mathevotaenia(Cestoda: Anoplocephalidae) and other tapeworms from marsupials in Argentina. Journal of Parasitology, vol. 89, pp. 1181–1185.Cardoso, TS, Costa-Neto, SF, Braga, C, Weksler, M, Simões, RO, Maldonado, A, Luque, JL & Gentile, R. 2020. Helminth metacommunity of small mammals in a Brazilian reserve: the contribution of environmental variables, host attributes and spatial variables in parasite species abundance. Community Ecology, vol. 21, pp. 159–170.Cardoso, TS, Simões, RO, Luque, JLF, Maldonado, A & Gentile, R., 2016. The influence of habitat fragmentation on helminth communities in rodent populations from a Brazilian Mountain Atlantic Forest. Journal of Helminthology, vol. 90, pp. 460–468.Costa-Neto, SF, Cardoso, TS, Boullosa, RG, Maldonado, A & Gentile, R. 2019. Metacommunity structure of the helminths of the black-eared opossum Didelphis auritain peri-urban, sylvatic and rural environments in south-eastern Brazil.Journal of Helminthology, vol. 93, pp. 720–731.D'Andrea, PS, Gentile, R, Maroja, LS, Fernandes, FA, Coura, R & Cerqueira, R. 2007. Small mammal populations of an agroecosystem in the Atlantic Forest domain, southeastern Brazil.Brazilian Journal of Biology, vol. 67, pp. 179-186.Dormann, CF, Gruber, B & Fruend, J. 2008. Introducing the bipartite Package: analysing ecological networks. R News, vol. 8, pp. 8–11.Geise, L. 2012. Akodon cursor (Rodentia: Cricetidae).Mammalian Species, vol. 44, pp. 33–43.Gentile, R, Cardoso, TS, Costa-Neto, SF, Teixeira, BR & D'Andrea, PS. 2018. Community structure and population dynamics of small mammals in an urban-sylvatic interface area in Rio de Janeiro, Brazil.Zoologia, vol. 35, pp. 1–12.Gibbons, LM. 2010. Keys to the Nematoda parasites of vertebrates: supplementary volume. CABI Publishing, London.Gibson, DI, Jones, A & Bray, RA. 2002. Keys to the Trematoda. CABI Publishing and the Natural History Museum, London, UK.Gomes, DC, Cruz, RP, Vicente, JJ & Pinto, RM. 2003. Nematode parasites of marsupials and small rodents from the Brazilian Atlantic Forest in the State of Rio de Neotropical Helminthology, 2022, 16(2), jul-dicGentile et al.art. 1=10-24art. 2=26-41Art 3 =42-51art. 4=52-59art. 5=60-65art 6=66-79art. 7=80-91art 8 =92-100art 9=102-111nota 1=112-116nota 2=118-123 rev =124-134
image/svg+xml171Janeiro, Brazil. Revista Brasileira de Zoologia, vol. 20, pp. 699–707. Gómez-Muñoz, MA, Robles, MR, Milano, AMF & Navone, GT. 2018. Helminth infection levels on Rattus rattus(Rodentia: Muridae) from Corrientes city, Argentina. Mastozoología Neotropical, vol. 25, pp. 221–227.Jiménez, FA, Catzeflis, F & Gardner, SL. 2011. Structure of parasite component communities of Didelphid marsupials: insights from a comparative study. Journal of Parasitology, vol. 97, pp. 779-786.Jones, A, Bray, RA & Gibson, DII. 2005. Keys to the Trematoda. CABI Publishing, U.K.Kajin, M, Cerqueira, R, Vieira, MV, Gentile, R. 2008. Nine-year demography of the black-eared opossum Didelphis aurita(Didelphimorphia: Didelphidae) using life tables. Revista Brasileira de Zoologia, vol. 25, pp. 206-213.Khalil, LF, Jones, A & Bray, RAA. 1994. Keys to the cestode parasites of vertebrates. CAB International, Wallingford, Oxon.Kruse, H, Kirkemo, AM & Handeland, K. 2004. Wildlife as source of zoonotic infections. Emerging Infectious Diseases, vol. 10, pp. 2067- 2072.Lafferty, KD & Kuris, AM. 1999. How environmental stress affects the impacts of parasites. Limnology & Oceanography, vol. 44, pp. 925–931.Lucio, CS, Gentile, R, Cardoso, TS, Oliveira-Santos, F, Teixeira, BR, Maldonado Jr., A & D'Andrea, PS. 2021. Composition and structure of the helminth community of rodents in matrix habitat areas of the Atlantic forest of southeastern Brazil. International Journal for Parasitology: Parasites & Wildlife, vol. 15, pp. 278–289.Maldonado, A, Gentile, R, Fernandes-Moraes, CC, D'Andrea, PS, Lanfredi, RM & Rey, L. 2006. Helminth communities of Nectomys squamipesnaturally infected by the exotic trematode Schistosoma mansoniin southeastern Brazil. Journal of Helminthology, vol. 80, pp. 369–375.Meerburg, BG, Singleton, GR & Kijlstra, A. 2009. Rodent-borne diseases and their risks for public health.Critical Reviews in Microbiology, vol. 35, pp. 221-270.Munõz, MAG, Robles, MDR, Milano, AMF & Navone, GT. 2018. Helminth infection levels on Rattus rattus(Rodentia: Muridae) from Corrientes city, Argentina. Mastozoología Neotropical, vol. 25, pp. 221-227.Panisse, G, Robles, MDR, Digiani, MC, Notarnicola, J, Galliari, C & Navone, GT. 2017. Description of the helminth communities of sympatric rodents (Muroidea: Cricetidae) from the Atlantic Forest in northeastern Argentina. Zootaxa, vol. 4337, pp. 243–262. Panti-May, JA, Palomo-Arjona, EE, Gurubel-González, YM, Barrientos-Medina, RC, Digiani, MC, Robles, MR, Hernández-Betancourt, SF & Machain-Williams, C. 2020. Patterns of helminth infections in Rattus rattusand Mus musculusfrom two Mayan communities in Mexico. Journal of Helminthology, vol. 94, e30.Pinto, RM, Kohn, A, Fernandes, BMM & Mello, DA. 1982. Nematodes of rodents in Brazil, with description of Aspidodera vicentei n.sp. Systematic Parasitology, vol. 4, pp. 263-267.Poulin, R. 2010. Network analysis shining light on parasite ecology and diversity. Trends in Parasitology, vol. 26, pp. 492–498.Santos-Rondon, MVS, Pires, MM, Reis, SF & Ueta, MT. 2012. Marmosa paraguayana(Marsupialia: Didelphidae) as a new host for Gracilioxyuris agilisis(Nematoda: Oxyuridae) in Brazil.Journal of Parasitology, vol. 98, pp. 170–174.Simões, RO, Souza, JGR, Maldonado, A & Luque, JL. 2011. Variation in the helminth community structure of three sympatric sigmodontine rodents from the coastal Atlantic Forest of Rio de Janeiro, Brazil.Journal of Helminthology, vol. 85, pp. 171–178.Travassos, L. 1937. Revisão da família Trichostrongylidea Leiper 1912. Memórias do Instituto Oswaldo Cruz, vol. 1, pp. 1–512.Vicente, JJ, Rodrigues, HO, Gomes, DC & Pinto, RM. 1997. Nematóides do Brasil. Parte V: nematóides de mamíferos.Revista Brasileira de Zoologia, vol. 14, pp. 1–452. Werk, DF, Gallas, M, Silveira, EF & Périco, E. 2016. New locality records for Guerrerostrongylus zetta(Travassos, 1937) Neotropical Helminthology, 2022, 16(2), jul-dicHelminths of small mammals in Rio de Janeiroart. 1=10-24art. 2=26-41Art 3 =42-51art. 4=52-59art. 5=60-65art 6=66-79art. 7=80-91art 8 =92-100art 9=102-111nota 1=112-116nota 2=118-123 rev =124-134
image/svg+xml172Sutton & Durette-Desset, 1991 (Nematoda: Heligmonellidae) parasitizing Oligoryzomys nigripes (Olfers, 1818) (Rodentia: Sigmodontinae) from southern Brazil. Check List, vol. 12, 1861.Yamaguti, S. 1961. Systema Helminthum. Interscience Publishers, New York.Received August 26, 2022.Accepted November 4, 2022.Neotropical Helminthology, 2022, 16(2), jul-dicGentile et al.art. 1=10-24art. 2=26-41Art 3 =42-51art. 4=52-59art. 5=60-65art 6=66-79art. 7=80-91art 8 =92-100art 9=102-111nota 1=112-116nota 2=118-123 rev =124-134