INTRODUCTION

Laelapidae mites (Mesostigmata, Dermanyssoidea) are distributed worldwide and are one of the most common ectoparasites of cricetid rodents (Strandtmann & Wharton 1958; Radovsky 1969). Knowledge of South American laelapids was primarily established by the pioneering work of Fonseca (1936, 1958), who described about 20 species, established the genera of some and defined the taxonomic limits of others. In South America, the study of laelapids was initiated by high rates of small mammal collecting due to a bubonic plague outbreak in north-eastern Brazil in the 1930s, and the creation of the National Plague Service (Servicio Nacional de Peste – SNP) (Fonseca 1958). Thousands of rodents were captured and the ectoparasites collected from the host pelage were systematically distributed among specialists for their study (Fonseca 1958). Laelapids were sent to the collections of the Museu de Zoologia da Universidade de Estado de São Paulo and Instituto Butantan in Brazil and were studied by Flavio Oliveira Ribeiro da Fonseca. This author reported (1958) that from the 4510 rodents captured in north-eastern Brazil, almost 60% were parasitized with mesostigmatid mites. Out of them, only 1500 rodents were identified to species level, and some of these species were reported in association with between 13-19 laelapid species such as Oligoryzomys nigripes (Olfers, 1818) (cited as O. eliurus in Fonseca (1958)).

The study of laelapids in Argentina began in the early 1960s with Ricardo A. Mauri from the Administración Nacional de Laboratorios e Institutos de Salud “Dr. Carlos G. Malbrán” (ANLIS) located in the Ciudad Autónoma de Buenos Aires in Argentina, when the mammologist Elio Massoia collected ectoparasites from their rodent hosts and delivered them to specialists for their study (Mauri 1965). It is worth noting that Massoia started to mention host-laelapid associations identified by Mauri in his publications, which was unusual for that period of time. The rodent O. nigripes was among the first to have parasitological information reported. Indeed, Massoia inaugurated by the term “eto-ecología” which seems self-evident nowadays and is also called “integrative-biology” (Massoia & Fornes 1964, 1972). Massoia himself did not emphasize the need for recognition of his work in epistemological terms (Contreras Roqué 2019). His comprehensive way of working (from collecting the remains of nests and excrements and identifying predators and parasites, to captive breeding of rodents at home for better biological observations) resulted in numerous and in many cases the only existent contributions of the ecology and ethology of small mammals (Pardiñas & Cirignoli 2000; Contreras Roqué 2019).

After the death of the mammalogist Abel Fornes in 1972, Massoia honored his memory with the description of a new species O. fornesiMassoia, 1973 from the genus Oligoryzomys Bangs, 1900. In fact, the genus Oligoryzomys deserves special interest in Massoia’s agenda and is still of current interest. Taxonomic uncertainties remain due to its genetic diversity and low morphological variation, including cryptic species (Weksler & Bonvicino 2015; Hurtado & D’Elía 2019).

Oligoryzomys nigripes is distributed in eastern Brazil, Paraguay, Uruguay and Argentina, and shares important areas of sympatry with O. flavescens (Waterhouse, 1837). In Argentina, O. nigripes reaches its southern limit at Reserva El Destino in Buenos Aires province (Udrizar et al. 2005), and O. flavescens extends southwards to the southwest of Buenos Aires province and the northern limit of Patagonia (Weksler & Bonvicino 2015). In addition, in the north-east of Argentina, O. flavescens shares geographical range in sympatry with O. fornesi and parapatry with O. nigripes concentrated in a sector of the Humid Chaco in Formosa and Chaco provinces (Weksler & Bonvicino 2015; Pardiñas 2017a).

From a parasitological point of view, the wide distribution of O. nigripes and O. flavescens and the sympatric and partly allopatric distribution of all three species constitute an interesting model to study host-parasite associations across their geographical range. The last two rodents share four mite genera: Androlaelaps, Gigantolaelaps, Mysolaelps and Laelaps (Fonseca 1958; Lareschi & Mauri 1998; Saraiva et al. 2012). For some of these mite genera, the degree of host specificity is debated (Gettinger 1987, 1992; Martins-Hatano et al. 2012; Silva-de La Fuente et al. 2020). While some laelapids are considered host-generalist parasites (Tipton et al. 1966; Furman 1972), others exhibit high host preference (e.g., Gettinger 1987, 1992; Martins-Hatano et al. 2012), with some of them species-specific to its sympatric hosts (Gettinger 1992; Lareschi & Galliari 2014). Few studies have analysed factors structuring host association of laelapids and rodent hosts in the Neotropics (e.g., Lareschi & Krasnov 2010; Linardi & Krasnov 2012). However, a comprehensive knowledge of the distribution of both host and mite is needed, especially when factors and patterns of distribution are studied to evaluate variation in the composition and structure of biological communities across space and/or over time (Krasnov et al. 2019).

Many cricetid rodent species and their laelapid associations have been identified and reported in Argentina, mainly for Buenos Aires province, while research in other areas of the country such as the northeast is scarce (see Lareschi & Mauri 1998). This is also the case for the genus Oligoryzomys, in particular because the previous surveys were conducted to explore the host-mite associations for all collected rodents rather than emphasizing a certain rodent genus. Consequently, this shows an uneven distribution of records for laelapid species associated with Oligoryzomys spp. In comparison with well-studied Buenos Aires province for the rodents O. nigripes and O. flavescens (Lareschi & Mauri 1998; Lareschi et al. 2003), only one locality record exists for the Entre Ríos province (Abba et al. 2001) and for Corrientes province (Lareschi et al. 2006b). There are scarce or no data of host-mite associations available where both rodents occur in sympatry, including Formosa and Chaco provinces. For other Oligoryzomys species from the same area, for example O. fornesi, the host-mite associations are yet unreported.

In this study we take the opportunity to honor Massoia’s academic legacy by presenting an updated account of laelapid mites associated with O. nigripes, O. fornesi and O. flavescens from samples collected over a wide geographical range in north-eastern and central Argentina over a decade.

MATERIAL AND METHODS

Mites were collected between 2007 and 2019 from O. nigripes, O. flavescens and O. fornesi from in 35 localities situated in Pampas, Espinal, Esteros del Iberá, Campos y Malezales, Chaco Húmedo, and Selva Parananense ecoregions (Burkart et al. 1999) located in six political provinces as detailed in Table 1 and in Fig. 1.

Rodents were identified by Ulyses Pardiñas (IDEAus) and Carlos Galliari (CEPAVE) and are housed at the Colección de Mamíferos del Centro Nacional Patagónico (CNP; Puerto Madryn, Chubut province, Argentina). Several specimens were used in genetic studies and their specific identifications were confirmed by molecular markers (Raúl González-Ittig, IDEA, unpublished data). We followed Teta et al. (2013); Pardiñas (2017a,b); Rivera et al. (2018), among others, for Oligoryzomys spp. distributed in north-western Argentina. Rodents from Jujuy, Tucumán and La Rioja provinces mentioned in the literature and reported as O. longicaudatus Bennett, 1832 and O. destructor Tschudi, 1844 (Autino et al. 2003) or O. f. occidentalis (López Berrizbeitia et al. 2013) are here recognized as O. brendae Massoia, 1998. We also followed Rivera et al. (2018) for the O. flavescens populations distributed in western Argentina and refer to them as O. f. occidentalis Contreras & Rosi, 1980. This was the case for species reported in Tucumán, La Rioja and Jujuy provinces (e.g., Mauri 1965).

Mites preserved in vials with 96% ethanol were cleared in lactophenol and mounted individually in Hoyer's medium for their identification in accordance with the original descriptions and drawings (Fonseca 1936, 1958) and keys provided by Tipton et al. (1966) and Furman (1971, 1972). Representative mites collected from every host species in each locality will be deposited at the Colección de Entomología del Museo de La Plata, Argentina (MLP). Laelapids have a holding number, which consists of the same field (e.g., CG, LTU, UP) or deposition acronym (e.g., CNP) as the hosts; for each individual mite of a single host a number was added and the letter M for mite, separated by a hyphen and MLP for Museo de la Plata (e.g., MLP-CNP6314/M3). A list of laelapid species is presented below with data about host species in each locality and number of specimens, all of them females. A brief report is also included with comments on the previous known geographical distribution and host associations with the emphasis on the genus Oligoryzomys.

RESULTS AND DISCUSSION

Subclass Acari Order Mesostigmata

Family Laelapidae Berlese, 1892 Subfamily Laelapinae Berlese, 1892 Genus Androlaelaps Berlese, 1903 Androlaelaps fahrenholzi (Berlese, 1911)

Type host: Unidentified host described as “Wild rat” Type-locality: Urbana, Illinois, USA

Material examined: 45 specimens.

Oligoryzomys flavescens: 12 specimens collected from BS1 (MLP-ROB128/M1, MLP-ROB130/M1), BS3 (MLP-CG/931/M1, MLP-CG930/M1), BS4 (MLP-CG688/M1, MLP-MK131/M1), BS6 (MLP- CNP6316/M1, MLP-LTU585/M1), CO1 (MLP-CNP5989/M1), ER1 (MLP-CG878/M1), ER2 (MLP-CG860/M1), and M12 (MLP-CG783/M1).

Oligoryzomys fornesi: 4 specimens collected from CH2 (MLP-CG404/M1), CH4 (MLP-CG408/M1, MLP-CNP5277/M1), and FO2 (MLP-LTU581/M1).

Oligoryzomys nigripes: 29 specimens collected from BS5 (MLP-CG919/M1, MLP-MK065/M1), CH1 (MLP-CNP1748/M1-M2,) CO4 (MLP- LTU422/M1-M2), CO5 (MLP-CNP5640/M1-M2), CO6 (MLP-CNP4737/M1, MLP-CNP5089/M1), FO1 (MLP-CG96/M1-M2), MI11 (MLP-CNP5725/M1- M2) MI2 (MLP-CG803/M1-M2), MI3 (MLP- CNP4281/M1-M2), MI4 (MLP-CNP6005/M1-M2), MI5 (MLP-CNP5634/M1), MI6 (MLP-CG476/M1- M2, MI7 (MLP-CNP4822/M1-M2), MI8 (MLP- CG602/M1-M2), and MI9 (MLP-CNP5973/M1-M2).

Remarks: Androlaelaps fahrenholzi presents a cosmopolitan distribution and a high variety of associations with rodents (Muridae, Cricetidae, Heteromyidae, Echimyidae, Sciuridae), marsupials (Didelphidae), and some birds (Strandtmann & Wharton 1958; Tipton et al. 1966; Furman 1972). Among Oligoryzomys species, A. fahrenholzi has been reported in association with O. nigripes in southern Brazil (Foz do Iguaçu) (Barros et al. 1993) and Uruguay (Lareschi et al. 2006b), and in Panama associated with Oligoryzomys fulvescens (Saussure, 1860) (Tipton et al. 1966). Also, in central and northern Argentina A. fahrenholzi was reported from a variety of mammals (Lareschi & Mauri 1998; Autino et al. 2003; Lareschi et al. 2003, 2019). O. flavescens and O. nigripes were reported as hosts of A. fahrenholzi in Buenos Aires and Entre Ríos provinces (Mauri 1965; Abba et al. 2001; Lareschi et al. 2003). The southern limit of A. fahrenholzi is located in Miramar associated with a variety of hosts including O. flavescens (Mauri 1965). Herein A. fahrenholzi is reported for the first time associated with O. flavescens and O. nigripes in the provinces of Corrientes, Misiones as well as in Formosa province with O. nigripes. In addition, O. fornesi is reported as a new host for A. fahrenholzi in Formosa, Corrientes and Chaco provinces. The new records extend the known distribution of the association O. nigripes - A. fahrenholzi from northeastern Buenos Aires province farther northeast to Corrientes and Misiones and similarly for O. flavescens - A. fahrenholzi including Entre Ríos province.

Additionally, the known southern limit of O. flavescens - A. fahrenholzi (Miramar) (Mauri 1965) is extended 460 km farther southeast to Sierra de la Ventana. This is the southern-most record for the species. Through the association with O. fornesi the known distribution of A. fahrenholzi in Argentina extends farther northwest for both O. nigripes and O. fornesi to Chaco and Formosa provinces.

Genus Gigantolaelaps Fonseca, 1939.

Gigantolaelaps wolffsohni (Oudemans, 1910)

Type-host: An unidentified rat

Type locality: Near Valparaíso, Chile. Material examined: 77 specimens.

Oligoryzomys flavescens: 30 specimens collected from BS1 (MLP-ROB10/M1-M2), BS2 (MLP-CNP6022/M1-M2), BS3 (MLP-CG930/M2-M3) BS4 (MLP-CG655/M1), BS5 (MLP-CG914/M1), BS6 (MLP-CNP6314/M6), CH1 (MLP-LTU585/M2), CO1 (MLP-CNP5960/M2-M3), CO3 (MLP-CNP5751/M1-M2), CO4 (MLP-CNP6004/M4-M5), CO5 (MLP-CNP5631/M1-M2), CO6 (MLP-CG383/M1-M2), ER1 (MLP-CG878/M2-M3), ER2 (MLP-CG860/M1-M2), ER3 (MLP-CNP5966/M1- M2), M1 (MLP-CNP5797/M1-M2), and MI2 (MLP-CG783/1M2-M3).

Oligoryzomys fornesi: 9 specimens collected from CH3 (MLP-CG404/M2), CH3 (MLP-CG404/M3), CH4 (MLP-CG408/M2), CO4 (MLP-CNP5617/M1- M2), FO1 (MLP-CNP5277/M2-M3), FO1 (MLP- CNP5277/M3), and FO2 (MLP-LTU581/M2-M3).

Oligoryzomys nigripes: 38 specimens collected from: BS5 (MLP-CG919/M2), CO1 (MLP- CNP5963/M1-M2), CO4 (MLP-LTU422/M3), CO5 (MLP-CNP4720/M1-M2), CO5 (MLP-CNP5640/M3- M4), CO6 (MLP-CNP5089/M2-M3), CO7 (MLP- CG609/M1-M2), CO8 (MLP-LTU409/M1-M2), CO9 (MLP-CNP5604/M1-M2), FO1 (MLP-CG96/M3- M4), MI1 (MLP-CG736/M1-M2), MI2 (MLP- CG803/M3-M4), MI3 (MLP-CNP5212/M1-M2), MI4 (MLP-LTU831/M1-M2), MI5 (MLP-CNP5800/M1- M2), MI6 (MLP-CNP4647/M1-M2), MI7 (MLP- CNP4822/M3-M4), MI8 (MLP-CG602/M3-M4), MI9 (MLP-CNP5973/M3-M4), and MI11 (MLP- CNP5974/M1-M2).

Remarks: Gigantolaelaps wolffsohni has a neotropical distribution and has been reported from Brazil, Venezuela, Chile (Lareschi & González-Acuña 2010), Uruguay (Lareschi et al. 2006b) and Panama (Tipton et al. 1966), mainly associated with oryzomyine species (Fonseca 1958; Furman 1972). Regarding the hosts of the present study in other countries, G. wolffsohni was reported in association with O. nigripes in north-eastern and southern Brazil (Gettinger 1987; Barros et al. 1993) and in association with O. nigripes and O. flavescens in Uruguay (Lareschi et al. 2006b). Also in Argentina, G. wolffsohni was registered in Buenos Aires province parasitizing O. flavescens and O. nigripes (Mauri 1965; Lareschi & Mauri 1998; Lareschi et al. 2003) with the southern limit in Miramar (Mauri 1965). Furthermore, the association between O. flavescens and G. wolffsohni was also reported in Villa Elisa (Entre Ríos province), San Juan Poriahú (Corrientes province) (Lareschi et al. 2006b) and Santa Inés (Misiones province) (Lareschi et al. 2019), where G. wolffsohni was reported parasitizing both O. flavescens and O. nigripes. Further associations with Oligoryzomys spp. for G. wolffsohni were reported for north-western Argentina in Tucumán province with O. brendae (Autino et al. 2003). Moreover, G. wolffsohni is reported from “rata silvestre” from San Pedro, Jujuy province (Mauri 1965; Lareschi & Mauri 1998). Also, in our study G. wolffsohni was collected from all three Oligoryzomys species, demonstrating a wide geographical distribution, and reflecting the distributional pattern of each host. Hence, we report the new association O. fornesi - G. wolffsohni for Chaco, Corrientes and Formosa provinces. Also this is the first report of the associations O. nigripes - G. wolffsohni for Corrientes and Formosa provinces, and O. flavescens - G. wolffsohni for Chaco province. Furthermore, the known southern limit in Argentina (Miramar) (Mauri 1965) of O. flavescens - G. wolffsohni is extended 460 km farther southeast to Sierra de la Ventana. The known northern limit of G. wolffsohni in Misiones province (Santa Inés) (Lareschi et al. 2019) is herein extended 300 km farther north for O. nigripes and O. flavescens. Additionally, new records in Corrientes province (associated with all three species) amplify the distribution comprehensively to the previously known locality San Juan Poriahú (Lareschi et al. 2006b). Thus, our study supports a broad geographical range of the association between O. nigripes, O. flavescens and G. wolffsohni from north-eastern Brazil, connecting with Uruguay and almost reaching the southern limit of each host’s distribution in Argentina. The association between O. fornesi and G. wolffsohni confirms the preference for the tribe Oryzomyini mentioned above (Gettinger 1987; Barros et al. 1993).

Genus Laelaps Koch, 1836

Laelaps paulistanensisFonseca, 1936

Type host: unidentified “wild rat” with the number N270

Type locality: São Paulo, Brazil Material examined: 43 specimens.

Oligoryzomys nigripes: 43 specimens collected from: BS5 MLP-CG919/M3), CH1 (MLP-CNP1748/M3- M4), CH2 (MLP-CNP6252/M1-M2), CO1 (MLP- CNP5963/M3-M3), CO4 (MLP-LTU422/M4-M5), CO5 (MLP-CNP5640/M5-M6), CO6 (MLP- CNP4737/M1), CO7 (MLP-CG608/M1-M2), CO8 (MLP-LTU409/M3-M4), CO9 (MLP-CNP6016/M1- M2), FO1 (MLP-CG96/M5-M6), MI1 (MLP- CG736/M4-M5), MI2 (MLP-CG803/M5-M6), MI3 (MLP-CNP5212/M3-M4), MI4 (MLP-LTU831/M3- M4), MI5 (MLP-CNP5800/M3-M4), MI6 (MLP- CNP4647/M1-M2), MI7 (MLP-CNP4822/M5- M6), MI8 (MLP-CG602/M5-M6), MI9 (MLP-CNP5973/M5-M6), MI10 (MLP-CNP6017/M1-M2), and MI11 (MLP-CNP5725/M3-M4).

Remarks: Laelaps paulistanensis has a wide neotropical distribution, with records from north-eastern to southern Brazil (Fonseca 1958; Nieri-Bastos 2004; Saraiva et al. 2012), Panama (Tipton et al. 1966), Venezuela (Furman 1972) and Uruguay (Lareschi et al. 2006b). In Argentina, L. paulistanensis was reported mainly associated with O. flavescens and O. nigripes, and was extensively recorded from both species in Buenos Aires province (Mauri 1965; Lareschi & Mauri 1998; Lareschi et al. 2003). In addition, L. paulistanensis was reported from both species in Villa Elisa (Entre Ríos province) (Abba et al. 2001), San Juan Poriahú (Corrientes province) (Lareschi et al. 2006b) and in Santa Inés (Misiones province) (Lareschi et al. 2019). In north-western Argentina, L. paulistanensis was reported associated with O. brendae in Tucumán province (Autino et al. 2003) and Euryoryzomys legatus (Thomas, 1925) in Jujuy province (cited as Oryzomys russatus in Autino et al. 2003). There is also a record of L. paulistanensis associated presumably with O. f. occidentalis in Tucumán province (Mauri 1965) and in La Rioja province associated with O. brendae (López Berrizbeitia et al. 2013).

In our study, L. paulistanensis is reported exclusively associated with O. nigripes. The reports herein presented extend the known distribution of L. paulistanensis in Argentina from Santa Inés (Misiones province) about 300 km farther north to Parque Provincial Urugua-í (MI8, Fig. 1, Table 1), and farther northwest to provinces of Chaco and Formosa, where the species is recorded for the first time. Also, for Corrientes province, the association O. nigripes L. paulistanensis in San Juan Poriahú (Lareschi et al. 2006b) is widely expanded with six new localities (Fig. 1, Table 1). Overall, the localities recorded in our study for L. paulistanensis collected from O. nigripes and the previous records for that host in Uruguay (Lareschi et al. 2006a) and Brazil (Fonseca 1958; Saraiva et al. 2012) reflect the distributional pattern of the host O. nigripes.

Laelaps schatziSavchenko & Lareschi, 2019 Type host: Oligoryzomys flavescens

Type locality: Pereyra, Partido de Berazategui, Buenos Aires, Argentina -34.837178°; -58.089833°

Material examined: 19 specimens.

Oligoryzomys flavescens: 11 specimens collected in BS3 (MLP-CG930/M4-M5), CH1 (MLP- LTU585/M3), CO5 (MLP-CN5606/M1-M2), ER1 (MLP-CG855/M1-M2), M1 (MLP-CG775/M3-M4), and MI2 (MLP-CG805/M1-M2).

Oligoryzomys fornesi: 8 specimens collected in CH1 (MLP-CNP5609/M1), CH3 (MLP-CG404/M4-M5), CH4 (MLP-CG408/M3), CO5 (MLP-CNP5626/M1-M2), and FO1 (MLP-CNP5010/M1-M2).

Remarks: Previously, L. schatzi has only been reported associated with its type host, O. flavescens, in north-eastern Buenos Aires province, Estancia Santa Ana de Carpinchori, in Entre Ríos province, Parque Provincial San Cayetano and Estancia San Juan Poriahú in Corrientes province, and at Estancia Santa Inés, Misiones province (Savchenko & Lareschi 2019). This rodent was previously reported associated with a morphologically similar mite species, L. paulistanensis (Lareschi et al. 2006a, 2019). In the course of this work, we reviewed specimens collected from O. flavescens and reported as L. paulistanensis in Santa Inés (Misiones province) (Lareschi et al. 2019) and subsequently identified them as the recently described species, L. schatzi. Furthermore, in the locality La Balandra (Buenos Aires province) where O. flavescens - L. paulistanensis associations were previously reported (Lareschi et al. 2003), we could identify the only O. flavescens - L. schatzi relationship. An extensive review of the O. flavescens L. paulistanensis association is needed.

Herein, we present a new host association between L. schatzi and O. fornesi, and in two new provinces, Chaco and Formosa. Also, L. schatzi is reported associated with O. fornesi in Corrientes province. Additionally, the association between O. flavescens and L. schatzi is newly reported for Chaco province. We also add further locality records for the association O. flavescens - L. schatzi for Entre Ríos and Misiones provinces. The habitat of both rodents is characterized by open vegetation and grasslands near water, and Massoia (1973) highlighted in the original description of O. fornesi its strong morphological resemblance to O. flavescens. Nevertheless, the taxonomic status of O. fornesi is still unclear, especially whether O. fornesi is a full (sister) species or a subspecies of O. flavescens and further morphological and cytogenetic data should be analyzed (Rivera et al. 2018). Only recently, O. fornesi was evaluated together with O. flavescens as part of the same species using unilocus delimitation methods and subsequently was considered as a junior subjective synonym of O. flavescens (Hurtado & D’Elía 2019). At least from the parasitological point of view the composition of laelapid species herein reported support the close relationship of the two hosts and do not exhibit characteristics to differentiate between them. Regardless of the actual specific status of O. fornesi further review of specimens of L. schatzi collected from O. fornesi is needed to explore potential differences at the intraspecific level for the nominal mite species between both rodents. Until now, the distribution of L. schatzi association for both hosts is restricted to Argentina. Especially for O. flavescens it will be interesting to explore whether the association includes the entire distribution of the host in other countries.

Genus MysolaelapsFonseca, 1936

Mysolaelaps microspinosusFonseca, 1936

Type-host. Undetermined rodent, captured with the host number (n°. 318)

Type-locality. São Paulo, Brazil. Material examined: 38 specimens.

Oligoryzomys flavescens: 22 specimens collected from BS1 (MLP-ROB61/M1-M2), BS2 (MLP-CNP6022/M3-M4), BS3 (MLP-CG930/M6-M7), BS4 (MLP-CG690/M5), BS5 (MLP-CG914/M2), BS6 (MLP-CNP6314/M7), ER3 (MLP-CNP5990/M3-M4), CO1 (MLP-CNP5989/M2-M3), CO2 (MLP- CNP5780/M1), CO5 (MLP-CNP5631/M3-M4), CO6 (MLP-CG383/M3-M4), M1 (MLP-CG770/M1-M2), and MI2 (MLP-CG805/M3-M4).

Oligoryzomys fornesi: 9 specimens collected from CH3 (MLP-CG413/M1-M2), CH4 (MLP-CG408/M4- M5), CO5 (MLP-CNP5626/M3-M4), FO1 (MLP-CNP5010/M3-M4), and FO2 (MLP-LTU581/M4).

Oligoryzomys nigripes: 7 specimens collected from BS5 (MLP-MK132/M1), CO6 (MLP-CG380/M1- M2), MI4 (MLP-CNP6008/M1-M2), and MI8 (MLP- CG532/M1-M2).

Remarks: The distribution of M. microspinosus was restricted to south-eastern Brazil after a broad survey, primarily associated with O. nigripes and O. mattogrossae (Fonseca 1958). Subsequent reports in Brazil did not expand the area of distribution but confirmed the association between O. nigripes and M. microspinosus (Lopes et al. 1989; Linardi et al. 1991). In central and southern Chile (Lareschi & González-Acuña 2010) M. microspinosus is reported in association with O. longicaudatus and in Uruguay associated with O. nigripes and O. flavescens (Lareschi et al. 2006b). This report supports the strong association with the genus Oligoryzomys, although the species is distributed north to Venezuela is associated with other Oryzomyines (Furman 1972). In central Argentina, M. microspinosus was extensively reported in Buenos Aires province associated with O. nigripes and O. flavescens (Lareschi & Mauri 1998; Lareschi et al. 2003) with the southern limit for the latter association in Miramar (Mauri 1965), and furthermore in Villa Elisa (Entre Ríos province) associated with O. flavescens and O. nigripes (Abba et al. 2001). Mysolaelaps microspinosus was also reported in Tucumán province associated with O. f. occidentalis (cited as O. flavescens in Mauri 1965) and O. brendae (Mauri 1965; Autino et al. 2003) and also associated with O. brendae in La Rioja province (López Berrizbeitia et al. 2013). Herein we report a new host of M. microspinosus, O. fornesi, in Corrientes, Chaco and Formosa provinces. The records for Chaco and Formosa provinces are reported for the first time. In addition, we report for the first time M. microspinosus associated with O. flavescens and O. nigripes in Corrientes and Misiones provinces. These reports extend about 300 km the known distribution of M. microspinosus in Argentina to the northernmost region in Misiones and in the northwest to Laguna Naineck (FO2, Fig. 1, Table 1) in Formosa province. Furthermore, M. microspinosus specimens from O. flavescens extend the known distribution of this mite in Argentina 460 km farther southeast to Sierra de la Ventana.

Mysolaelaps parvispinosusFonseca, 1936

Type-host. Type species was described from two cotypes captured on a wild rat of undetermined species (°n. 226) and topotypes of rats (°ns. 269 and 270).

Type-locality. São Paulo, Brazil Material examined: 44 specimens.

Oligoryzomys flavescens: 5 specimens collected from CO1 (MLP-CNP5960/M4), CO4 (MLP-CNP6004/M6), CO5 (MLP-CNP5631/M5), MI1 (MLP-CG751/M1), MI2 (MLP-CG805/M5).

Oligoryzomys fornesi: 9 specimens collected from CH3 (MLP-CG404/M6-M7), CO1 (MLP- CNP5962/M1), CO5 (MLP-CNP5626/M5-M6), FO1 (MLP-CG446/M1-M2), FO2 (MLP-LTU581/M5-M6).

Oligoryzomys nigripes: 30 specimens collected from CO4 (MLP-LTU422/M6-M7), CO6 (MLP- CNP4737/M2-M3), CO8 (MLP-LTU409/M5- M6), FO1 (MLP-CNP5084/M1-M2), MI1 (MLP- CG736/M6-M7), MI2 (MLP-CG803/M7-M8), MI3 (MLP-CNP5212/M5-M6), MI4 (MLP-CNP6025/M1- M2), MI5 (MLP-CNP5800/M5-M6), MI6 (MLP-CG591/M1-M2), MI7 (MLP-CG600/M1-M2), MI8 (MLP-CG602/M7-M8), MI9 (MLP-CNP5973/M7- M8), MI10 (MLP-CNP6017/M3-M4), and MI11 (MLP-CNP5974/M3-M4).

Remarks: The known distribution of M. parvispinosus reaches from northeast to south Brazil (Fonseca 1958; Nieri-Bastos 2004; Cruz et al. 2012), northern Peru (Lareschi & Velazco 2013), Venezuela (Furman 1972) and Panama (Tipton et al. 1966), primarily associated with Oligoryzomys spp. and some other oryzomyine species. In Brazil, one of the main reported hosts is O. nigripes (e.g., Fonseca 1958; Botelho & Williams 1980; Linardi et al. 1991; Barros et al. 1993). In Venezuela, M. parvispinosus was reported associated with several species from the Tribe Oryzomyini, mainly O. fulvescens (Furman 1972), and in Peru collected from O. destructor (Lareschi & Velazco 2013). In Argentina, M. parvispinosus was registered for the first time in Misiones province associated with Oryzomys sp. (Mauri 1982). Subsequently, M. parvispinosus was reported only two other times, in San Juan Poriahú, Corrientes province (Lareschi et al. 2006b) associated with O. flavescens and in Estancia Santa Inés, Misiones province (Lareschi et al. 2019) associated with O. flavescens and O. nigripes.

Herein, the association between O. fornesi and M. parvispinosus is reported for the first time, as well as the record of M. parvispinosus in Formosa and Chaco provinces. Considering the previous records for other Oligoryzomys species, these reports are not unexpected. Also, the known association with O. nigripes was extended to Formosa province. Furthermore, the report of M. parvispinosus with O. nigripes extends the known distribution of this association from Santa Inés (Misiones province) to about 300 km farther north and adds three new localities for Corrientes province. Therefore, in Argentina the new northern limit of the O. nigripes M. parvispinosus relationship is Parque Provincial Urugua-í (MI8, Fig. 1, Table 1), and in the south in Paso de los Libres (CO6, Fig. 1, Table 1). This locality in Corrientes province is now the most southern report of the species M. parvispinosus (in this case associated with O. nigripes). Additionally, for Corrientes province, the known distribution of the association O. flavescens - L. paulistanensis in San Juan Poriahú (Lareschi et al. 2006b) is amplified with three new localities (Fig. 1, Table 1). Hence, M. parvispinosus broadly reflects the distribution of O. nigripes in Brazil, reaching Uruguay and Argentina. It is unknown why the species is not collected farther south in Entre Ríos and Buenos Aires provinces in the Pampas and Espinal ecoregions (Burkart et al. 1999), respectively, although the corresponding hosts are present there. Also in the north of Corrientes province, M. parvispinosus is encountered in various ecoregions including Delta e Islas del Paraná but is absent in the same ecoregion farther south of the province. Several biotic or abiotic factors (Krasnov et al. 2019) may play an important role in the distribution of M. parvispinosus but additional studies are needed to determine them.

CONCLUSION

The results obtained in this study present new host and locality records for laelapids associated with species of Oligoryzomys from north-eastern and central Argentina. This paper presents the first records of all mentioned records for Formosa and Chaco provinces, as well as the host-mite associations for the rodent O. fornesi. Previous reports of mites associated with O. fornesi can be found for central Brazil (Gettinger 1987, 1992); however the taxonomy of the host species has been revised and most populations are now referred to O. mattogrossae (Allen, 1916) (see Weksler et al. 2017). Thus, these references from Brazil were not considered as host-mite records for the host O. fornesi and were interpreted as O. mattogrossae. Nevertheless, the host-mite associations presented for O. fornesi, distributed in Argentina, are reported for the first time.

The distribution of previously reported associations between O. flavescens and O. nigripes and corresponding laelapids is expanded to new provinces including Misiones, and new localities such as Corrientes province are now included. The three species of Oligoryzomys reported in the study were parasitized with A. fahrenholzi, G. wolffsohni, M. microspinosus and M. parvispinosus. Laealps paulistanensis was associated exclusively with O. nigripes, whereas L. schatzi was associated only with O. flavescens and O. fornesi. These mites have wide distributions in Argentina, some extending more than 1 500 km from Sierra de la Ventana to Parque Provincial Urugua-í, covering several of those ecoregions described by Burkart et al. (1999). A partial exception is M. parvispinosus, unrecorded in Espinal and Pampas ecoregions (in Buenos Aires and Entre Ríos provinces, respectively) but represented in Delta e Islas del Paraná although limited to the northern region of Corrientes province. Perhaps M. parvispinosus has a comparatively higher dependency to the environment than the other laelapid species. The decreasing gradient in temperature and humidity, from warm, subtropical climate in the north to temperate pampean conditions in the south may affect the development of the species. In any case, further studies are necessary to confirm this hypothesis.

Combined integrative work between mammalogists and parasitologists is needed and highly recommended to assure correct identification of both the host and the mite. Broad surveys of host-mite associations have enormous value for many biological questions (co-evolution, parasitism, factors of distributional patterns, host-specificity, etc.) as well as zoonotic studies. For instance, O. nigripes and O. flavescens are important as hosts of a group of viruses of the family Hantaviridae (Carbajo & Teta 2009; González-Ittig et al. 2014) which cause the Hantavirus Pulmonary Syndrome, a disease of high mortality in humans (Martinez et al. 2010). Oligoryzomys flavescens can also be associated with the bacterium Leptospira interrogans, responsible for Leptospirosis, a zoonotic systemic disease (Lovera et al. 2017).

The role of mesostigmatid mites in the circulation of some disease agents has lately received more attention globally, and is being attributed mainly to the members belonging to the family Laelapidae (Miťková et al. 2015). Especially the genus Laelaps has been shown to harbour rickettsial agents (Kuo et al. 2020). The list of host-mite associations presented here is an incentive to continue the research, as it raised many interesting questions to address in the future.

Acknowledgments

We are especially grateful to Carlos Galliari (CEPAVE) and Ulyses Pardiñas (IDEAus, CONICET, Argentina) for host identification, and to Raúl González-Ittig (IDEA) for specific species identifications supported by molecular data; to C. Galliari, María del Rosario Robles, Guillermo Panisse, Jorge Barneche, Natalia Guerreiro, María Cecilia Ezquiaga, Agustín Abba, Luis Giambelluca, U. Pardiñas, Juliana Notarnicola , Erika Cuellar, Juliana Sanchez and Mara Urdapilleta for field assistance and to Natália Tóthová (Universtity College London, U.K.) for English revision. The authors also thank an anonymous reviewer for his comments that contributed significantly to our study. Field work was supported by the Agencia Nacional de Promoción Científica y Tecnológica (PICT924 to Graciela Navone, and PICT 2015-1564 to M. Lareschi). Studies in the laboratory were supported by the Agencia Nacional de Promoción Científica y Tecnológica (PICT 2015-1564) and the Universidad Nacional de La Plata, Argentina (both to M. Lareschi). Collecting permits were provided by the Ministerio de Agroindustria de la Provincia de Buenos Aires, Ministerio de Ecología, Recursos Naturales Renovables y Turismo de la Provincia de Misiones, Ministerio de la Producción y Ambiente de la Provincia de Formosa, Dirección de Recursos Naturales de la Provincia de Corrientes, and Dirección de Fauna de la Provincia del Chaco. This study is part of the doctoral dissertation of E. Savchenko at the Facultad de Ciencias Naturales y Museo, Universidad Nacional de La Plata, Argentina.

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Información adicional

Editor asociado: U. F. J. Pardiñas & C. A. Galliari

Cite as: Savchenko, E., M. Melis & M. Lareschi. 2021. Laelapid mites (Mesostigmata) ectoparasites of Oligoryzomys (Rodentia: Cricetidae) in north-eastern and central Argentina. Mastozoología Neotropical, 28(1):e0526. https://doi.org/10.31687/saremMN.21.28.1.0.05