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Necromys Benefactus Populations in Central Argentina

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Necromys Benefactus Populations in Central Argentina Mastozoología Neotropical, 12(2):261-268, Mendoza, 2005 ISSN 0327-9383 ©SAREM, 2005 Versión on-line ISSN 1666-0536 MORPHOMETRIC AND ALLOZYMIC CHARACTERIZATION OF NECROMYS BENEFACTUS POPULATIONS IN CENTRAL ARGENTINA María C. Provensal1, Gladys E. Calderón2, Marina B. Chiappero3, Cristina N. Gardenal3, Jaime J. Polop1, and Marta S. Sabattini2 1 Departamento de Ciencias Naturales, Universidad Nacional de Río Cuarto, Río Cuarto, Córdoba, Argentina. <[email protected]>. 2 Instituto Nacional de Enfermedades Virales Humanas, Pergamino, Argentina. 3 Cátedra de Genética de Poblaciones y Evolución, Fac. Cs. Exactas, Físicas y Naturales, Universidad Nacional de Córdoba, Córdoba, Argentina. Key words. Genetic distance. Necromys. Morphometric variation. Oliveros virus. Pampa virus. The genus Necromys Ameghino, 1889 (Ro- Aires province and southern Uruguay and N. dentia, Sigmodontinae) is polytypic and wide- benefactus would have a broader distribution, spread in central-eastern South America, com- from the south-east intermountain area of prising several different species distributed in Buenos Aires province to the western and the Central Andes, the Chacoan and Pampean northeastern Buenos Aires, eastern of La regions and the northeastern Argentina, east- Pampa, south of Santa Fe, and north of Córdoba ern and southern Brazil, and Uruguay (Reig, province. No further information is available 1987; Galliari and Pardiñas, 2000). about distinctive morphometric and/or genetic Massoia and Fornes (1967) recognized two characters useful to properly classify field speci- different species in the Pampean region and mens of these two species of Necromys. Uruguay; these authors argued that populations It is well known that rodents are the most from southern Uruguay and southeastern important natural reservoirs for zoonotic dis- Buenos Aires province correspond to Necromys eases (Hugh-Jones et al., 1995), although only obscurus and those in the north-west of Buenos a small proportion of the organisms infecting Aires province belong to N. benefactus. Reig rodents would cause disease in humans. In the (1987) proposed that populations from Uru- decade of 1990, a new arenavirus called guay and southern Santa Fe, Córdoba, north- Oliveros (OLV) was isolated from rodents ern La Pampa, and northwestern Buenos Aires captured at Oliveros and J.B. Molina (south- belong to the species N. obscurus, considering ern Santa Fe province), identified in the field N. benefactus as a subspecies; those popula- as Bolomys obscurus (Mills et al., 1996) and tions from the south-east and south-west of then called N. benefactus (Galliari and Pardiñas, Buenos Aires province would represent a new 2000). In 1992, a genomic variant of OLV species. Results of morphological and morpho- denominated Pampa (PAM) was isolated from metric analysis by Galliari and Pardiñas (2000) individuals captured at Maciel (southern Santa reinforce the taxonomic and distributional hy- Fe province), apparently of the same rodent pothesis of Massoia and Fornes (1967) for the species. Serologic evidence of PAM infection Pampean species of Necromys: N. obscurus in the rodent reservoir has been reported in would be restricted to southeastern Buenos several localities of the central region of Ar- Recibido 4 febrero 2005. Aceptación final 24 junio 2005. 262 Mastozoología Neotropical, 12(2):261-268, Mendoza, 2005 M. C. Provensal et al. www.cricyt.edu.ar/mn.htm gentina (Riera, 2004). The pathogenesis of determination was made then on these indi- OLV virus is not known to date, but the emer- viduals, on the basis of qualitative cranial char- gence of this new infectious agent poses the acters described by Galliari and Pardiñas need to improve our knowledge on biological (2000): nasals flat, nasofrontal suture with re- properties of the host populations. spect to the orbital process, frontal edges, in- The precise determination of species bound- terorbital constriction, free upper border of zy- aries and relationships among taxa in gomatic plate, shape of the mesopterygoid Sigmodontinae rodents is particularly difficult fossa, and direction of the coronoid process in in some groups, mainly because of the poor relation to articular process and develop of the morphological differentiation accompanying the coronoid process. speciation process in several genera (Steppan, One hundred and forty four specimens of 1993). In this paper, we characterize and com- ³60 days old from the first generation of both pare individuals assigned to the genus colonies were compared separately by sex, Necromys trapped in different localities of the considering the results of a previous study of Pampean region of Argentina, using morpho- morphometric characters in animals of differ- metric variables and allozymic frequencies. ent age classes (Polop et al., 2000). The ani- The specimens of the genus Necromys were mals were sacrificed by inhalation of anesthetic obtained at three localities: Oliveros (32°34’S, gas (Methoxyflurane), and the following char- 60°51’W) and Uranga (33°16’S, 60°42’W) in acters measured: body weight (BW), nasal Santa Fe province and Pergamino (33°32’S, width (NW), length of hard palate (PL), length 60°49’W) in Buenos Aires province (Fig. 1). of upper diastema (D), mandible length (ML), Individuals were captured in the weedy bor- zygomatic width (ZW), and nasal length (NL). ders of crop fields and roads with Sherman Body weight was recorded to the nearest 0.5 g live traps set in lines of 125 m long, with one in animals 60 to 120 days old, and the arith- trap placed every 5 m. A preliminary taxo- metic mean for each group (according to sex nomic determination of Necromys individuals and location) was compared. Measurements was made in the field on the basis of external were taken with caliper and recorded to the features (size, fur coat and coloration) and total, nearest 0.02 mm. tail, ear and right hind foot lengths. Twelve The similarity between colonies and between specimens from Uranga (5 males and 7 females) adults from Uranga and Pergamino were es- and 47 from Pergamino (22 males and 25 fe- tablished using the D2 of Mahalanobis males) were carried alive to a field laboratory (Mahalanobis, 1936; Rao, 1952). A cluster of the National Institute of Human Viral Dis- analysis was performed by the UPGMA pro- eases (Pergamino, Argentina) in order to es- cedure, based on pairwise Euclidean distances tablish two laboratory colonies. The animals among individuals from all colonies and sites were maintained under controlled conditions (male and female individuals analyzed sepa- (18°-22°C; 12L: 12D); food and water were rately) provided ad libitum. Comparative morphomet- Allozymic studies: In order to examine the ric studies were performed in these specimens degree of similarity in the gene pool of the and their descendants. populations assigned to the species of genus Additional animals from Uranga (N = 11) Necromys, we analysed the allozymic polymor- and Pergamino (N = 20) and specimens phism in individuals captured in Oliveros trapped in Oliveros (N = 24) were sacrificed (N=24), Uranga (N=11) and Pergamino in the field, liver and kidneys were removed (N=20). Twelve informative loci from 9 enzy- and preserved in liquid nitrogen for allozymic matic proteins were studied: lactate dehydro- analyses. genase (Ldh, E.C. 1.1.1.27), leucine aminopep- Morphological studies: Specimens captured tidase (Lap, E.C. 3.4.11), malic enzyme (ME, in Uranga and Pergamino were sacrificed after E.C. 1.1.1.40), a-glycerophosphate dehydro- they left descendants. A second taxonomic genase (Gpdh, E.C. 1.1.1.8), esterases (Est-5 CHARACTERIZATION OF NECROMYS BENEFACTUS POPULATIONS 263 Fig. 1. Capture localities of Necromys specimens in the central-eastern Argentina. and Est-6, E.C. 3.1.1.1), kidney and liver acid and f value will be significant and positive. If phosphatases (AcpK-1, AcpK-2 and AcpL, E.C. each sample comes from a single panmictic 3.1.3.2), glutamate oxaloacetate transaminase population, f will not be statistically different (Got-1, E.C. 2.6.1.1), glucose-6-phosphate from 0. dehydrogenase (G6pdh, E.C. 1.1.1.49) and 6- Mean heterozygosity per locus, percentage phosphogluconate dehydrogenase (6-PGD, E.C. of polymorphic loci (P95% criterion) and mean 1.1.1.44). Preparation of homogenates, elec- number of alleles per locus (A) were calcu- trophoresis and staining to reveal enzyme ac- lated using BIOSYS-2 (Black, 1997). tivity was performed as described by Selander Specimens captured in the different locali- et al. (1971), and Gardenal and Blanco (1985). ties presented the following characteristics with Deviations from Hardy-Weinberg equilibrium respect to the fur coat and coloration: the dor- in each locus in each population were tested sum had short guard hairs (9-10 mm), with a by the exact test based on a Markov chain yellowish pheomelanin portion, giving the back algorithm (Guo and Thompson, 1992) using an agouti colour. The sides of the body were GENEPOP version 3.2a (Raymond and yellowish-grey and the abdomen was whitish Rousset, 2000). Genetic structure of popula- grey, with white gular and inguinal regions. tions was estimated by F-statistics, using the The range of the length of head and body was unbiased estimators of Weir and Cockerham 135-209 mm and the range of the length of tail q (1984): for FST and f for FIS. Significance of was 48-84 mm. These features correspond with q and f values were obtained by randomization the description of Necromys benefactus made procedures, as implemented in FSTAT program by Galliari and Pardiñas (2000). (Goudet, 2000). If there are two cryptic spe- Considering the morphological studies all the cies, each population sample will contain indi- specimens captured in Uranga, Pergamino and viduals pertaining to two separate gene pools those obtained from the laboratory colonies 264 Mastozoología Neotropical, 12(2):261-268, Mendoza, 2005 M. C. Provensal et al. www.cricyt.edu.ar/mn.htm showed the following states of qualitative char- served genotypic frequencies did not deviate acters: a) nasal projected rostrally, overhang- from those expected under Hardy Weinberg ing the premaxillae and hiding the incisors in equilibrium.
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