Seasonal shifts within juvenile recruit sex ratio of Pampas mice (Akodon azarae)

Sex ratio (SR) variation of Akodon azarae juvenile recruits was analysed during 1985–86 on a Pampean grassland in central Argentina. Rodents were intensively live-trapped on a 0.81 ha grid. The SR (proportion of males) of the overall population did not differ significantly from evenness (0.49). In contrast, juvenile recruits (0.5–2.0 months of age) showed SR fluctuations according to their time of birth. SR was strongly female-biased among spring and autumn juveniles (0.13 and 0.33, respectively), while males were predominant (0.72) among the summer juveniles.
Social and demographic implications of these results are discussed in the framework of current SR theory. Spring female recruits should be selected for summer reproduction and autumn female recruits for winter survival and spring reproduction. The overwintered population was characterized by 1:1 SR and an age structure composed of males older than females. Most of these males had been recruited as juveniles during the summer. They also made up most of the resident male population in the spring. A selective allocation of energy by the mothers among their offspring is the mechanism proposed to explain the seasonal changes in SR, as pregnant females which gave birth to the summer recruits were in better physical condition than the mothers of both spring and autumn recruits. These results are consistent with the predictions of Werren & Charnov (1978), based on temporal overproduction of one or the other sex according to differential changes in the life-history expectations for both sexes.     

A new species of Syphacia (Nematoda: Oxyuridae) from Akodon azarae (Rodentia: Cricetidae) in Argentina

Eight species of Syphacia (Nematoda: Oxyuridae) have been reported from South America in rodents of the Sigmodontinae, only 1 of which has been recorded in Argentina. Syphacia (Syphacia) carlitosi n. sp. is described from the ceca of Akodon azarae bibianae and Akodon azarae hunteri (Sigmodontinae: Akodontini) captured in 3 provinces in the northeast region of Argentina. The new species is differentiated principally by the shape of the cephalic plate; distribution of submedian papillae and amphids; presence, extent, and shape of cervical alae in females; absence of lateral alae; absence of deirids; spicular and gubernaculum length; shape and structure of accessory hook of gubernaculum; and distance of mamelons, excretory pore, and vulva from the anterior extremity. This is the second record of Syphacia parasitizing rodents of the tribe Akodontini.     

Meiotic behavior of gonosomically variant females of Akodon azarae (Rodentia, Cricetidae)

The meiotic behavior of sex chromosomes has been investigated in variant females of Akodon azarae, both in pachytene oocytes and metaphase I. In somatic cells, these females have a heteromorphic sex pair, in which the minor chromosome has been previously interpreted as a major deletion of the long arm of the X chromosome (dX). After microspreading for synaptonemal complex analysis, pachytene oocytes show two axes of very different lengths (100:17.1), which correspond to the sex chromosomes X and dX. True synapsis is abnormally restricted (43.3%) between these sex chromosomes; on the other hand, self-synapsis of both the X and dX chromosomes is frequent (60%). Single, nonsynapsed axes or axial segments are thickened. Strong chromatin condensation occurs around nonsynapsed axes or axial segments, giving many of these sex pairs an appearance similar to an XY body ("sex vesicle"). The minor gonosome axis differs from that of the Y chromosome of male meiosis, as the former is shorter (relative to the X) and has a different synaptic behavior. In 17 metaphases I from XdX variant females, only heteromorphic, end-to-end joined sex pairs were observed. These variant females differ from the variant females of the wood lemming Myopus schisticolor in several respects, but a similar mechanism seems to be prevalent in other species of the genus Akodon. Self-synapsis of unequal gonosomes in oocytes is assumed as an escape from functional deterioration, following the hypothesis put forward by others.     

Chromosomal aspects and inheritance of the XY female condition in Akodon azarae (Rodentia,Sigmodontinae)

The populations of several species of Akodon present, besides XX females, a variable proportion of fertile XY females. In Akodon azarae, a correspondence exists between the X-chromosome C-banding pattern and the sexual phenotype of XY individuals: males carry a determinate X-chromosome type, defined by its C-banding pattern, and XY females, any of two others. To confirm the relation between X-chromosome type and the XY female condition and to investigate the hereditary transmission of these different X-chromosomes, we analyzed 50 animals captured in the field and 95 individuals corresponding to the F₁ and F₂ offspring of 16 crosses.It was seen that the correlation between X type and the sexual phenotype of XY animals is retained, and that the three X types are transmitted to the progeny. It was also observed that the male offspring of XY females receive the X-chromosome from their male parents and the Y from their mothers. These results strongly support the causal role of an X-borne mutation in A. azarae XY sex reversal, and discard a mutation of the Y-chromosome as the sole basis of this phenomenon.     

A new angiostrongylid (Nematoda) species from the pulmonary arteries of Akodon azarae (Rodentia: Cricetidae) in Argentina

Angiostrongylus morerai n. sp. (Nematoda: Angiostrongylidae) is described from the pulmonary arteries of Azara's grass mouse Akodon azarae (Rodentia: Cricetidae) in Argentina. It is distinguished from its congeners principally by the morphology of the dorsal ray, which is as long, or longer, than the externodorsals and has 2 long branches; the spicule lengths are also greater (400-465 microm). This is the first record of a metastrongyloid from sigmodontine rodents in Argentina.     

The sex-determining zinc finger sequences in XY females of Akodon azarae (Rodentia, Cricetidae)

Wild populations of Akodon azarae comprise females with a karyotype indistinguishable from that of males. These individuals were formerly assumed to be Xx, the x being an X chromosome with a deletion of most of its long arm. By using a DNA probe derived from the testis-determining region of the human Y chromosome (comprising a candidate gene for the testis-determining factor, Y-linked zinc finger [ZFY]), we demonstrate that A. azarae gonosomally variant females are XY and not Xx. The ZFY sequences in A. azarae are amplified and located in two different families of EcoRI fragments derived from Y-chromosome DNA. No rearrangement or change in the state of methylation of ZFY or ZFX (X-linked zinc finger) sequences were found in XY females. We propose that sex reversal in A. azarae may be mediated by a gene or genes other than ZFX or ZFY.     

Phenotypic flexibility of digestive morphology and physiology of the South American omnivorous rodent Akodon azarae (Rodentia: Sigmodontinae)

We have determined the occurrence of responses at different levels (morphological, physiological and biochemical) in the omnivorous rodent Akodon azarae upon cold acclimation (15 degrees C). A short-term enhancement in food consumption appeared to account for the maintenance of both mass and body composition. At the morphological level, the main response was an increase in the dimensions of small intestine, which constitutes the section of the gut where absorption and secretion take place. An increase in sucrase specific activity was only found in small intestine. Sucrose independent maltase activity was very low since 99.8% of total maltase activity was due to sucrase-isomaltase (SI) complex. Protease specific activities were not affected. The fact that resting metabolic rates determined at 15 and 23 degrees C were similar in cold acclimated animals suggests a change in lower critical temperature. In conclusion, our results show that A. azarae exhibits different strategies to support cold environment that could lead to an enhancement in digestion and absorption efficiency. Furthermore, this work suggests that low temperature is an independent cue of other environmental factors to trigger the strategies allowing the maintenance of body condition in A. azarae.     

Inter-male aggression with regard to polygynous mating system in Pampean grassland mouse, Akodon azarae (Cricetidae: Sigmodontinae)

Based on the hypothesis that, in Akodon azarae, polygyny operates through female defence, we studied inter-male aggression in order to test the following predictions: during the breeding period (1) resident males are more aggressive than intruder males in the presence of females (FP), and (2) aggressive behaviour is independent of male condition (resident or intruder) in the absence of females (FA). To test our predictions, we used the resident male behavioural response towards an intruder male in relation to FP or FA. We conducted 30 encounters in FP and 27 in FA in 0.79-m² round enclosures placed in the Espinal Reservation. Our results support the prediction that, in FP, the intensity of aggressive behaviour exhibited by males varied in relation to resident or intruder condition. Resident males showed high levels of aggression towards intruders, and intruders exhibited the greatest values of submissive behaviours with residents. In FA, the intensity of aggressive behaviour did not vary in relation to resident or intruder condition. Both resident and intruder males exhibited low aggressive behaviour and inter-male encounters resulted mainly in non interactive behaviours. Our results support the hypothesis that, in A. azarae, the polygynous mating system operates through female defence.     

Phenotypic plasticity in response to low quality diet in the South American omnivorous rodent Akodon azarae (Rodentia: Sigmodontinae)

We studied the responses in the omnivorous rodent A. azarae submitted to a low quality diet at morphological, physiological and biochemical levels. At short term, a decrease in body mass occurred. A later increase in food consumption constituted a strategy that allowed a temporal recovery of physical condition. However, hyperphagia appeared not to be enough to maintain physical condition after 30 days of low quality diet consumption. At the morphological level, an increase in length (9%) of the anterior portion of the gut occurred, the part of the gut where digestion and absorption take place. A decrease in small intestine weight could be related with the long-term impairment of body condition. Inhibition of sucrase specific activity in small intestine would indicate a down-regulation of sucrase-isomaltase complex. Total maltase specific activity in small intestine was not affected suggesting an up-regulation of sucrase-independent maltase specific activity. A down-regulation of protease specific activity in small intestine occurred in response to low quality diet. The specific activity of disaccharidases in caecum and large intestine was down-regulated. The strategies and constraints at different levels of A. azarae upon low quality diet are discussed.     

Effects of high density on spacing behaviour and reproduction in Akodon azarae: A fencing experiment

We studied the short term spacing behavioural responses of Pampean grassland mouse (Akodon azarae) with regard to population density in four 0.25 ha enclosures (two control and two experimental) in the 2011 breeding season. Based on the hypothesis that A. azarae breeding females exhibit spacing behaviour, and breeding males show a fusion spatial response, we tested the following predictions: (1) home range size and intrasexual overlap degree of females are independent of population density values; (2) at high population density, home range size of males decreases and the intrasexual home range overlap degree increases. To determine if female reproductive success decreases at high population density, we analyzed pregnancy rate, size and weight of litters, and period until fecundation in both low and high enclosure population density. We found that both males and females varied their home range size in relation to population density. Although male home ranges were always bigger than those of females in populations with high density, home range sizes of both sexes decreased. Females kept exclusive home ranges independent of density values meanwhile males decreased home range overlap in high breeding density populations. Although females produced litters of similar size in both treatments, weight of litter, pregnant rate and period until fecundation varied in relation to population density. Our results did not support the hypothesis that at high density females of A. azarae exhibit spacing behaviour neither that males exhibit a fusion spatial response.     

Bot fly parasitism (Rogenhofera bonaerensis) (Diptera, Cuterebridae) in the pampean grassland mouse (Akodon azarae), in Argentina

Seasonality and impact of parasitism by the larvae of Rogenhofera bonaerensis (Diptera: Cuterebridae) in pampean grassland mouse (Akodon azarae, Rodentia, Cricetidae) populations were studied in grasslands and cropfield borders near Diego Gaynor (Buenos Aires Province, Argentina), from November 1985 to December 1986. Trapped mice infections ranged from one larva (77%) to four (4%) per host; larvae occurred on tails (75.7%), backs (17.1%), thighs (5.7%), and abdomen (1.4%). Prevalence of larvae occurred in late spring (November and December) (35%), and in fall (March to June) (20%), with a peak of 25% during April and May. In spring these larvae affected 46% of resident and 19% of transient individuals; however, no differences were found between residents and transients in spring or fall. There was no significant relationship between mouse population density and prevalence of parasitism (r = -0.56). The annual percentage of infected rodents changed with age (body size classes): adults (37%) greater than subadults (25%) greater than juveniles (16%); with no differences between the sexes. Similar results were recorded in the spring, but there were no differences in the fall. Significant differences were not detected between stable (22% infected) and disturbed habitats (14% infected). During winter uninfected mice survived better than those infected during the previous fall. In contrast, the summer survival among spring infected mice and noninfected mice was similar. Parasitism showed no effects on reproductive activity and on home range size. Larvae showed strong host specificity for A. azarae (99%). Only one case was recorded in the long-tailed mouse (Oligoryzomys flavescens), and none in the white paunch mouse (Calomys laucha), both species sympatric with A. azarae.(ABSTRACT TRUNCATED AT 250 WORDS)     

A new species of Liniscus (Nematoda: Trichuridae) from Oxymycterus rufus and Akodon azarae (Cricetidae: Sigmodontinae) in Buenos Aires Province, Argentina

Liniscus diazae n. sp. (Nematoda: Trichuridae) is described from the urinary bladder of Oxymycterus rufus and Akodon azarae (Cricetidae: Sigmodontinae) from Buenos Aires Province, Argentina. Liniscus diazae can be differentiated from its congeners by having a much longer spicule and without an elevated vulva. In addition, L. diazae can be distinguished from L. incrassatus and L. himizu by having a longer body length. Males are similar in posterior width to L. papillosus and L. himizu, but thinner than L. incrassatus and L. maseri. This is the second record of a capillarid from mammals in Argentina. An updated list of capillarid species of rodents and insectivores from North America, with their synonyms, hosts, sites of infection, and geographic distribution, is provided.     

Diversity of the helminth community of the Pampean grassland mouse (Akodon azarae) on poultry farms in central Argentina

This paper describes the helminth community of the Pampean grassland mouse (Akodon azarae) inhabiting poultry farms in central Argentina. Winter diversity (season of high rodent abundance) has been compared to spring diversity (season of low rodent abundance). Species richness was seven in both seasons: five nematodes (Syphacia carlitosi, Stilestrongylus spp., Trichuris laevitestis, Pterygodermatites (Paucipectines) azarai and Protospirura numidica criceticola) and two cestodes (adults of Cyclophyllidea and Taenia taeniaeformis hepatic cysts). No difference in richness was detected between host sexes in each season or among host age classes. However, the helminth community showed 67% similarity between winter and spring, with diversity being significantly higher in spring (H = 0.873) than in winter (H = 0.546; P < 0.0005). This could be attributed to different factors, such as host abundance, host diet or environmental factors, that affect the transmission of each species differently. On the other hand, Stilestrongylus spp. and S. carlitosi showed higher dominance and intensity in both seasons compared to their cohabiting species, P. (P.) azarai and T. laevitestis, respectively. The lower values of the latter two species may be related to a crowding effect due to their large body sizes. This is the first report of cestodes in A. azarae. The finding of T. taeniaeformis strobilocerci could be important in the epidemiology of parasitosis in domestic animals of the farms.     

Return to preferred habitats (edges) as a function of distance in Akodon azarae (Rodentia, Muridae) in cropfield-edge systems of central Argentina

We studied the ability of Akodon azarae (Rodentia, Muridae) to return to their preferred habitat, when released at a perpendicular distance (25, 50 or 75 m) from the edge towards the cropfield, and a parallel distance (100 m) away from the site of first capture within the edge habitat. Return success was estimated as the proportion of animals recovered in edges. The recapture rate between the field and the border was significantly higher than the recapture rate estimated according to successive captures in the border. Successful returns did not decrease significantly with increasing release distance, but animals released at 50 m from edges were less successful in returning to borders than the other release-distance groups. Although the median time taken to the first recapture in edges did not differ among the release-distance groups, rodents released at 25 m and 50 m returned to edges faster than those released at 75 m. A. azarae showed both a successful return to the edge and a trend to return to the home range area. We conclude that A. azarae can return to edges from cropfields at distances that are larger than those they usually travel, allowing the use of fields when they present good conditions for reproduction and survival. Successful return is probably the result of direct movements rather than random wandering.     

Multiple origins of XY female mice (genus Akodon): phylogenetic and chromosomal evidence

Despite the diversity in sex determination across organisms, theory predicts that the evolution of XY females is rare in mammals due to fitness consequences associated with infertility or the loss of YY zygotes. We investigated this hypothesis from a phylogenetic perspective by examining the inter- and intraspecific distribution of Y chromosomes in males and females (XY females) in South American field mice (Akodon). We found that XY females occurred at appreciable frequencies (10-66%) in at least eight Akodon species, raising the possibility that this system of sex determination has arisen multiple times independently. To determine the number of origins of XY females in Akodon, we constructed a molecular phylogeny of 16 species of Akodon based on mitochondrial DNA control region sequences. Both parsimony and maximum-likelihood reconstruction of ancestral states suggest that multiple steps (gains or losses of XY females) best explain the evolution of XY females, but do not clearly differentiate between single and multiple origins. We then directly compared functional and non-functional Y chromosomes in six species by Southern blot analysis. We found that male and female Y chromosome restriction fragment length polymorphism patterns were identical within species, but always differed between species, providing evidence that XY females arose at least six times within the Akodon lineage. To our knowledge, this pattern in Akodon is the first documentation of a novel sex-determining system arising multiple times within a tight clade of mammals. In addition, this system provides a clear test of the accuracy of phylogenetic methods to reconstruct ancestral states.     

Density-dependent habitat selection between maize cropfields and their borders in two rodent species (Akodon azarae and Calomys laucha) of Pampean agroecosystems

We studied habitat preferences and intra and interspecific density-dependent effects on habitat selection by Akodon azarae and Calomys laucha between maize fields and their adjacent borders, during different developmental stages of the crop. Akodon azarae detected quantitative differences between habitats, using preferentially borders throughout the year, while C. laucha perceived borders and cropfields as quantitatively similar during spring and summer and it detected borders as quantitatively better at the high density period (autumn and winter). These results support the prediction of differential habitat preferences as a model of community organisation at the low density period, while they are consistent with shared habitat preferences during autumn and winter when both species apparently coexist in the better habitat (border). Akodon azarae showed intraspecific density-dependent habitat selection throughout the year, except in spring, while habitat selection by C. laucha was density-dependent in spring, autumn and winter. The effect of interspecific density on habitat selection was detected in both habitats and changed seasonally. The effect of A. azarae over C. laucha by resources exploitation was detected in borders, while competitive effects of C. laucha over A. azarae was observed within cropfields. Both species were more affected by exploitation competition than interference, which was more common in borders than in maize fields. We conclude that seasonally have a profound effect in habitat selection of these species because it changes the intensity of intra and interspecific competition and affects different habitat preferences and basic suitability of habitats. This revised version was published online in November 2006 with corrections to the Cover Date.     

Ectoparasite occurrence associated with males and females of wild rodents Oligoryzomys flavescens (Waterhouse) and Akodon azarae (Fischer) (Rodentia: Cricetidae: Sigmodontinae) in the Punta Lara wetlands, Argentina

The objective of this research was to study infestation parameters and indexes of ectoparasites associated with each sex of the wild rodents Oligoryzomys flavescens (Waterhouse) andAkodon azarae (Fischer) in the Punta Lara wetlands, Argentina. A trend towards higher mean abundance (MA) and ectoparasite specific richness was observed in males of O. flavescens whereas those values were similar for both A. azarae sexes. The prevalence of the following ectoparasites was significantly higher on males (P<0.05): Mysolaelaps microspinosus Fonseca (65.2%) and Hoplopleura travassosi Werneck (73.9%) on O. flavescens, and Ixodes loricatus Neumann (71.4%) on A. azarae. Only H. travassosi mean abundance was significantly higher on males (MA=44.1). Since I. loricatus and Hoplopleura spp. are involved in the transmission of pathogens that cause diseases in animals and humans, and whose reservoirs are rodent hosts, these results are epidemiologically important.