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Mastozoología neotropical

versão impressa ISSN 0327-9383versão On-line ISSN 1666-0536

Mastozool. neotrop. vol.26 no.2 Mendoza jun. 2019  Epub 14-Dez-2019

 

NOTAS

ACTIVITY AND GROUP-LIVING IN THE PORTER’S ROCK RATS, Aconaemys porteri

Patrón de actividad y comportamiento social de Aconaemys porteri

María José Frugone1 

Loreto A Correa2  3 

Raúl Sobrero4  5 

1Instituto de Ecología y Biodiversidad, Departamento de Ciencias Ecológicas, Facultad de Ciencias, Universidad de Chile

2Escuela de Medicina Veterinaria, Facultad de Ciencias. Universidad Mayor

3Departamento de Ecología, Facultad de Ciencias, Pontificia Universidad Católica de Chile

4Laboratorio de Ecología de Enfermedades, Instituto de Ciencias Veterinarias del Litoral (ICiVet-Litoral), Universidad Nacional del Litoral (UNL)

5Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET)

Abstract

We provide the first systematic data on behavior and ecology of Aconaemys porteri. We used telemetry to monitor patterns of activity, resting locations, and range areas. Rock rat movements were statistically similar during nighttime and daytime, implying no clear diurnal or nocturnal activity. Animals used from 2 to 9 putative resting locations, but one was used more frequently. Rock rats showed relatively smaller range areas and low-to-moderate spatial overlap with neighbors, compared to other rodent species. These results indicate that A. porteri exhibit an intermediate level of sociality, compared to other octodontids.

Palabras clave ámbito de hogar; patrón de actividad; Octodontidae; vida en grupo

Resumen

Proporcionamos los primeros datos obtenidos de manera sistematizada sobre el comportamiento y la ecología de Aconaemys porteri. Utilizamos telemetría para monitorear los patrones de actividad, sitios de descanso o refugios, y ámbitos de hogar. Los movimientos de A. porteri tanto para las horas de oscuridad como de luz fueron estadísticamente similares, sugiriendo un patrón de actividad no estrictamente diurno o nocturno. Los animales utilizaron desde 2 hasta 9 sitios de descanso siendo uno de ellos utilizado en mayor frecuencia. Aconaemys porteri exhibió ámbitos de hogar relativamente pequeños y escaso a moderado solapamiento espacial entre individuos. Estos resultados sugieren que A. porteri presenta un nivel de sociabilidad intermedio, en comparación con otros octodóntidos.

Palabras clave ámbito de hogar; patrón de actividad; Octodontidae; vida en grupo

Among caviomorph rodents, social organization ranges from solitary living to social forms, in which individuals interact frequently, overlap their range areas, and share resting locations (e.g. Ebensperger et al. 2004; Ebensperger & Hayes 2008). These rodents include species that differ morphologically and physiologically and use a great diversity of habitats (Verzi et al. 2015) which seems to covary with groupliving (Lacey & Ebensperger 2007).

Thus, caviomorphs offer unique opportunities to examine factors associated with variation in social organization across and within species (e.g. Maher & Burger 2011). Within caviomorphs, the octodontids (Octodontidae) comprises 16 extant species grouped in six living genera with surface-dwelling (Octomys), fossorial (Octodontomys, Octodon, Tympanoctomys), completely subterranean (Spalacopus) and semisubterranean (Aconaemys) habits (Ojeda et al. 1996; Gallardo et al. 2007; Lessa et al. 2008).

Aconaemys or rock rats used open areas and forests, on both eastern and western slopes of the Andes (Pearson 1984; Gallardo & Mondaca 2002).

Three species are currently recognized: A. fuscus, A. sagei, and A. porteri (Gallardo & Mondaca 2002). Aconaemys fuscus inhabits highland forests of Araucaria araucana (Araucariaceae) and sandy areas (Muñoz-Pedreros 2000), while A. sagei inhabits ungrazed bunchgrass and Nothofagus forest with bamboo cover (Pearson 1984). Aconaemys porteri is distributed from Volcán Villarrica to Puyehue in Chile, and from both Parque Nacional Lanín and Parque Nacional Nahuel Huapi in Argentina (Pearson 1984; 1995; Gallardo & Reise 1992; Gallardo & Mondaca 2002). In these areas, Porter’s rock rats inhabit dense bamboo and southern beech rainforests (Pearson 1983; 1984; Gallardo & Reise 1992).

Evaluating the extent of the social behavior of Aconaemys remains critical to determine whether ecological conditions and species-specific attributes drove the evolution of group living in the octodontids. A recent comparative analysis suggested that social behavior in caviomorphs was gained and lost repeatedly, perhaps originating from an ancestral species that was socially flexible, and where the loss of group-living has been associated to the use of habitats with high plant cover (Sobrero et al. 2014a). Moreover, robust knowledge about multiple ecological factors (distribution of resources such as food, predation risk, and soil conditions associated with digging burrows or nesting sites) surely will contribute to build a theory of sociobiology that is closer and more consistent with the diversity of mammalian social behavior (Tang-Martínez 2003; Hayes et al. 2007). Taken together, establishing how group-living varies across species is critical for comparative studies to examine the origin and the adaptive value of this behavior (Blumstein & Armitage 1998; Ebensperger 2001; Ebensperger & Blumstein 2006). The scarce available and anecdotal evidence suggests that Porter’s rock rats lives in small groups in communal burrow systems and both diurnal and nocturnal activity has been recorded (Pearson 1983; Verzi et al. 2015). Also, in captivity, individuals displayed a high tolerance for conspecifics (Verzi et al. 2015); however it has not been tested in natural populations and laboratory conditions can modify behavior of captive individuals (Calisi & Bentley 2009).

Our aim in this research note is to report data of the activity patterns and social behavior of freeliving A. porteri. The study site was located at the Fundo San Pablo de Tregua (39°35’S, 72°05’W), Panguipulli, Chile.

San Pablo de Tregua is characterized by a dry and short summer where mean annual precipitation is around 5000 mm, and ambient temperature ranges from 5°C to 20°C (Alvarez & Lara 2008; Guevara et al. 2015). The site consisted of Valdivian rainforest where dominant tree species were coigüe (Nothofagus dombeyi), raulí (N. alpina), and tepa (Laureliopsis philippiana) (Schlegel & Donoso 2008). Animals were captured using a combination of 172 (14 x 14 x 40-cm) Tomahawk (model 201; Tomahawk Live Trap Company, Hazelhurst, Wisconsin) and leg-hold traps during 8 days in January, 2011. Qualitatively, putative nest sites used by A. porteri are structurally similar to burrows and resting locations of O. degus (Fulk 1976; Lessa et al. 2008), consisting mainly of oblique tunnels connecting the surface to the nest, with several branches and active openings (4-5) associated to patches of perennial bamboo (Chusquea quila), covering part of the burrow system. We defined rats burrow as areas of 1.5 m2 covered by vegetation, in which we found signs of A. porteri presence (i.e., feces) and where radiocollared individuals were found during trapping and telemetry periods. Ground mounds were sometimes found outside openings and fresh feces in burrows openings allow us to determine if each burrow system was active.

Moreover, in two occasions rats’ vocalizations were listened (but not recorded and analyzed) during trapping. We placed traps near burrow openings and inside patches with high bamboo cover and baited them with rolled oats, cereals and sunflower seeds. Traps were opened 08:00h and closed 00:00h and checked every hour. During each capture, we recorded sex, body mass and reproductive status, and each animal was marked with an ear tag (Monel 10051; National Band and Tag Co., Newport, Ky., USA). Low number of rats represents the species at a normal density, based on knowledge about population ecology of other caviomorph rodents (e.g. Cassini 1991; Ebensperger et al. 2008). All adult-sized individuals (N = 5) were fitted with a radiocollar weighing 7–9 g (BR radiocollars; AVM Instrument Co., Colfax, California) with unique pulse frequencies. Weight of radio-collars represented about 4–5% of body adult weight (e.g. Ebensperger et al. 2008). At the end of our study all radiocollared animals were recaptured and radiocollars were removed (Ebensperger et al. 2004; 2012).

During 5 days and nights it was performed homing technique every two hours, to determine resting locations or putative nest places, using LA 12-Q receiver (for radiocollars tuned to 150.000–151.999 MHz frequency; AVM Instrument Co., Colfax, California) and a handheld 3-element yagi antenna (AVM instrument Co., Colfax, California). Once located, the position of each animal was marked with flagging material coded for individuals. Each radiofix location was then referenced twice with a Garmin portable global positioning system (Garmin International Inc., Olathe, Kansas), precision always was within 5 m. The determination of group composition required the compilation of a symmetric similarity matrix of pairwise association of the resting locations of all adult animals during homing (Whitehead 2008). Social organization was quantified based on the number and sex composition of adult members in a social group (Ebensperger & Hayes 2016). Thus, we conducted a hierarchical cluster analysis of the association matrix in SOCPROG software (Whitehead 2009).

To estimate daily activity patterns and range areas, we recorded locations hourly of all radiocollared animals for 5 days and 4 nights in 2011, at nighttime (21:00-07:00 h) and daytime (07:00-21:00 h). Sunrise occurred at approximately 06:30 h, whereas sunset occurred at 20:30 h. The spatial location of animals was determined using triangulation (Kenward 2001). We used 2 LA 12-Q receivers, each connected to a null peak antenna system (AVM Instrument Co.). Every null peak system had four 7-element yagi antennas. Distance between antenna stations was about 85 m. Bearings from both antenna stations were then transformed into x–y locations with the software Locate II (Nams 1990). We calculated the distance traveled (in m), between successive scans as a measure of aboveground activity of the Porter’s rock rats. The same individuals were monitored throughout consecutive days and nights. As a result, locations recorded at 24-h intervals were not independent of one another. Consequently, and for statistical analyses, we divided the entire data collection period into 5 days and 4-night cycles, defined on the basis of sunrise and sunset at study site. For the daytime portion of each activity cycle, we calculated the mean distance travelled for each radiocollared individual. We used a similar approach for the nighttime portion of the activity cycle. As a result, each radiocollared individual contributed 2 dependent data points to our analysis of activity. We used repeated-measures analysis of variance to examine the effect of activity time (day versus night) on individual activity of males and females.

The range area was determined from locations recorded through triangulation and included animal resting locations. While triangulation is thought to interfere less with the activity of radio-collared animals compared with homing (Kenward 2001; Ebensperger & Blumstein 2006), the topography and cover type of our study site precluded the use of longrange radio-fixings because of signal bounce. Thus, although we used the homing technique, we previously trained ourselves to locate animals quickly to minimize disrupting their navigation or locomotion behavior (see Ebensperger et al. 2008). Data points from each individual were mapped using the 95% minimum convex polygon algorithm in Ranges 6 (Kenward et al. 2003). Pairwise estimates of the percent overlap between polygons for different animals also were calculated using Ranges 6. We compared the mean size (in m2) of range areas and percent range overlap by male and female Porter’s rock rats with Mann–Whitney U-tests. We used Wilcoxon matched-pair tests to compare percent overlap in range areas of individuals assigned to the same burrow location associations and percent overlap that these individuals had with individuals assigned to different associations in January 2011. All statistical analyses were calculated using Statistica 7.0 (StatSoft Inc. 1984–2004) and results are reported as mean ± SE.

Animals used 2 to 9 burrow locations, namely where one or more rats were found repeatedly during total radioscans (n=22). We recorded 21 scans in which animals shared burrow or resting locations. Of these, 20 observations involved male–female pairs, and 1 involved 2 males–1 female associations. Distance moved between radioscans was variable through time of day or night (Fig. 1). The distance moved between any two consecutive radio fixes during the night averaged 49 ± 28 m (n = 5, range: 1-249 m), and was 1.4 times larger than distance moved during day (36 ± 25 m, range: 0.5-425 m). However, this difference was not statistically significant (F8,9 = 0.420, P = 0.535). Five individuals were radio-tracked during the day and night, which provided an average of 98 useful radio fixes per animal. Spatial overlap among radio-collared Porter’s rock rats was relatively low (20 ± 5%) ranging from 3% to 34% (Fig. 2). If data from all individuals are combined, the size of range areas averaged 48 ± 13 m2 (n = 5). When sex was examined, females tended to range over larger areas (51 ± 3 m2, n = 3) than males (45 ± 39 m2, n = 2), a non-statistically significant difference (Mann–Whitney U test, z = 3.0, p > 0.10).

Fig. 1 Mean (± 1 SE) distance moved (m) since previous scan of Porter’s rock rats (Aconaemys porteri) monitored every 1 h for 4 days and 6 nights at San Pablo de Tregua. 

Fig. 2 Range areas (95% minimum convex polygons) of five Porter’s rock rats (Aconaemys porteri) during the activity period. The arrow signals the geographic north. 

Taken together, and based on this short-term study, results suggest that A. porteri are social. However, compared to other octodontids, social behavior of A. porteri seems relatively low, in terms of their low spatial overlap and group size. For example, evidence for Aconaemys suggests that A. fuscus forms social group up to 7 individuals (Muñoz-Pedreros 2000). On the other hand, similar studies, using telemetry techniques in O. degus, revealed range areas over 1 ha in summer with spatial overlap between individuals ranging from 12.5-59.4% (Quirici & et al. 2010). Our results suggest a smaller group size for A. porteri, however future studies are recommended, including larger sample sizes or different populations, to confirm results reported here. Additionally, our results support that A. porteri is active during daytime and nighttime. Continuous pattern of activity in other octodontids (Octodon lunatus) may be the consequence of relatively stable microclimatic or cover conditions (Jensen et al. 2003; Sobrero et al. 2014b). Plant cover has been shown to be an important factor for the distribution (e.g. Birney et al. 1976), social group size and space use in caviomorphs, included octodontids (Quirici & et al. 2010; Sobrero et al. 2014b) and cavies (Asher et al. 2004; Taraborelli 2008). Compared with range areas of other octodontids like Octodon degus (Hayes et al. 2007), O. lunatus (Sobrero et al. 2014b), and Octodontomys gliroides (Rivera et al. 2014), A. porteri showed small range areas, where polygons matched the spatial distribution of bamboo patches. Finally, these results are consistent with the hypothesis of the evolution of sociality in Octodontids, in which social living would be a derive characteristic evolving relatively recently from solitary-living ancestors. This novel evidence gives new insights into the social behavior of this species and the evolution of sociality across caviomorph rodents.

Acknowledgments

We are very thankful to Daniela Mellado, Gabriel Ortega, Carmen Paz Silva and Freddy Mondaca for field assistance. We are indebted to Juan Avila (Fundo San Pablo de Tregua, Universidad Austral de Chile) for providing all necessary facilities to access the field sites. Special thanks to Mauricio Soto-Gamboa and Luis Ebensperger for the economic support, equipment, and for their critical comments on previous versions of the manuscript. This study was supported by FONDECYT (grant 10900302 to Luis Ebensperger and grant 3150306 to RS).

REFERENCIAS

B01 Alvarez P. C., & A. Lara. 2008. Crecimiento de una plantación joven en fajas con especies nativas en la Cordillera de Los Andes de la provincia de Valdivia. Bosque (Valdivia) 29:181-191. https://doi.org/10.4067/s0717-92002008000300001Links ]

B02 Asher, M., E. S. De Oliveira, & N. Sachser. 2004. Social system and spatial organization of wild guinea pigs (Cavia aperea) in a natural population. Journal of Mammalogy 85:788-796. https://doi.org/10.1644/bns-012Links ]

B03 Birney, E. C., W. E. Grant, & D. D. Baird. 1976. Importance of vegetative cover to cycles of Microtuspopulations. Ecology 57:1043-1051. https://doi.org/10.2307/1941069Links ]

B04 Blumstein, D. T., & K. B. Armitage. 1998. Life history consequences of social complexity: a comparative study of grounddwelling sciurids. Behavioral Ecology 9:8-19. https://doi.org/10.1093/beheco/9.1.8Links ]

B05 Calisi, R. M., & G. E. Bentley. 2009. Lab and field experiments: Are they the same animal? Hormones and Behavior 56:1-10. https://doi.org/10.1016/j.yhbeh.2009.02.010Links ]

B06 Cassini, M. H. 1991. Foraging under predation risk in the wild guinea pig Cavia aperea. Oikos 62:20-24. https://doi.org/10.2307/3545441Links ]

B07 Ebensperger, L. A. 2001. A review of the evolutionary causes of rodent group-living. Acta Theriologica 46:115-144. [ Links ]

B08 Ebensperger, L. A., & D. T. Blumstein. 2006. Sociality in New World hystricognath rodents is linked to predators and burrow digging. Behavioral Ecology 17:410-418. https://doi.org/10.1093/beheco/arj048Links ]

B09 Ebensperger, L. A., & L. D. Hayes. 2008. On the dynamics of rodent social groups. Behavioural Processes 79:85-92. https://doi.org/10.1016/j.beproc.2008.05.006Links ]

B10 Ebensperger, L. A., R. Sobrero, V. Campos, & S. M. Giannoni. 2008. Activity, range areas, and nesting patterns in the viscacha rat, Octomys mimax. Journal of Arid Environments 72:1174-1183. https://doi.org/10.1016/j.jaridenv.2008.02.003Links ]

B11 Ebensperger, L. A., M. J. Hurtado, M. Soto-gamboa, E. A. Lacey, & A. T. Chang. 2004. Communal nesting and kinship in degus (Octodon degus). Naturwissenschaften 91:391-395. https://doi.org/10.1007/s00114-004-0545-5Links ]

B12 Ebensperger, L. A. et al. 2012. Ecological drivers of group living in two populations of the communally rearing rodent, Octodon degus. Behavioral Ecology and Sociobiology 66:261-274. https://doi.org/10.1007/s00265-011-1274-3Links ]

B13 Ebensperger, L. A., & L. D. Hayes. 2016. Causes and evolution of group-living. Sociobiology of Caviomorph Rodents: An Integrative Approach (L. A. Ebensperger & L. D. Hayes, eds.). John Wiley & Sons. https://doi.org/10.1002/9781118846506.ch7Links ]

B14 Fulk, G. W. 1976. Notes on activity, reproduction, and socialbehavior of Octodon degus. Journal of Mammalogy 57:495-505. https://doi.org/10.2307/1379298Links ]

B15 Gallardo, M. H., & F. Mondaca. 2002. The systematics of Aconaemys (Rodentia, Octodontidae) and the distribution of A. sagei in Chile. Mammalian Biology 67:105-112. https://doi.org/10.1078/1616-5047-00015Links ]

B16 Gallardo, M. H., R. Ojeda, C. A. Gonzalez, & C. A. Rios. 2007. The Octodontidae revisited. The quintessential naturalist: honoring the life and legacy of Oliver P. Pearson (D. A. Kelt, E. P. Lessa, J. Salazar-Bravo & J. L. Patton, eds.). University of California Publications in Zoology. [ Links ]

B17 Gallardo, M. H., & D. Reise. 1992. Systematics of Aconaemys (Rodentia, Octodontidae). Journal of Mammalogy 73:779-788. https://doi.org/10.2307/1382195Links ]

B18 Guevara, G., R. Godoy, P. Boeckx, C. Jara, & C. Oyarzún. 2015. Effects of riparian forest management on Chilean mountain in-stream characteristics. Ecohydrology & Hydrobiology 15:160-170. https://doi.org/10.1016/j.ecohyd.2015.07.003Links ]

B19 Hayes, L. D., A. S. Chesh, & L. A. Ebensperger. 2007. Ecological predictors of range areas and use of burrow systems in the diurnal rodent, Octodon degus. Ethology113:155-165. https://doi.org/10.1111/j.1439-0310.2006.01305.xLinks ]

B20 Hayes, L. D., & L. A. Ebensperger. 2011. Caviomorph rodent social systems: an introduction. Journal of Mammalogy 92:1-2. https://doi.org/10.1644/10-mamm-s-187.1Links ]

B21 Jensen, S. P., S. J. Gray, & J. L. Hurst. 2003. How does habitat structure affect activity and use of space among house mice? Animal Behaviour 66:239-250. https://doi.org/10.1006/anbe.2003.2184Links ]

B22 Kenward, R. E. 2001. A Manual for Wildlife Radio Tagging. Academic Press, San Diego, CA, USA. [ Links ]

B23 Kenward, R. E., A. B. South, & S. S. Walls. 2003. Ranges 6, version 1.2: for the analysis of tracking and location data., Wareham, United Kingdom. [ Links ]

B24 Lacey, E. A., & L. A. Ebensperger. 2007. Social structure in octodontid and ctenomyid rodents. Rodent societies: an ecological and evolutionary perspective (J. O. Wolff & P. W. Sherman, eds.). University of Chicago Press, Chicago, Illinois, USA. [ Links ]

B25 Lessa, E. P., A. I. Vassallo, D. H. Verzi, & M. S. Mora. 2008. Evolution of morphological adaptations for digging in living and extinct ctenomyid and octodontid rodents. Biological Journal of the Linnean Society 95:267-283. https://doi.org/10.1111/j.1095-8312.2008.01057.xLinks ]

B26 Maher, C. R., & J. R. Burger. 2011. Intraspecific variation in space use, group size, and mating systems of caviomorph rodents. Journal of Mammalogy 92:54-64. https://doi.org/10.1644/09-mamm-s-317.1Links ]

B27 Muñoz, A., & R. Murúa. 1987. Biología de Octodon bridgesi bridgesi (Rodentia, Octodontidae) en la zona costera de Chile central. Boletín de la Sociedad de Biología de Concepción (Chile) 58:107-117. [ Links ]

B28 Muñoz-Pedreros, A. 2000. Orden Rodentia. Mamíferos de Chile (A. Muñoz-Pedreros & J. Yáñez-Valenzuela, eds.). CEA Ediciones, Valdivia. [ Links ]

B29 Nams, V. O. 1990. Locate II user’s guide. Pacer Computer Software, Tatamagouche, Nova Scotia, Canada. [ Links ]

B30 Ojeda, R. A., J. M. Gonnet, C. E. Borghi, S. M. Giannoni, C. M. Campos, & G. B. Díaz. 1996. Ecological observations of the red vizchacha rat, Tynpanoctomys barrerae in desert habitats of Argentina. Mastozoología Neotropical 3:183-191. [ Links ]

B31 Pearson, O. P. 1983. Characteristics of a mammalian fauna from forests in Patagonia, southern Argentina. Journal of Mammalogy 64:476-492. https://doi.org/10.2307/1380360Links ]

B32 Pearson, O. P. 1984. Taxonomy and natural-history of some fossorial rodents of Patagonia, southern Argentina. Journal of Zoology 202:225-237. https://doi.org/10.1111/j.1469-7998.1984.tb05952.xLinks ]

B33 Pearson, O. P. 1995. Annotated keys for identifying small mammals living in or near Nahuel Huapi National Park or Lanin National Park, southern Argentina. Mastozoología Neotropical 2:99-148. [ Links ]

B34 Quirici, V. et al. 2010. Seasonal variation in the range areas of the diurnal rodent Octodon degus. Journal of Mammalogy 91:458-466. [ Links ]

B35 Rivera, D. S., S. Abades, F. D. Alfaro, & L. A. Ebensperger. 2014. Sociality of Octodontomys gliroides and other Octodontid rodents reflects the influence of phylogeny. Journal of Mammalogy 95:968-980. https://doi.org/10.1644/14-mamm-a-057Links ]

B36 Schlegel, B. C., & P. J. Donoso. 2008. Effects of forest type and stand structure on coarse woody debris in old-growth rainforests in the Valdivian Andes, south-central Chile. Forest Ecology and Management 255:1906-1914. https://doi.org/10.1016/j.foreco.2007.12.013Links ]

B37 Sobrero, R., O. Inostroza-Michael, C. E. Hernandez, & L. A. Ebensperger. 2014a. Phylogeny modulates the effects of ecological conditions on group living across hystricognath rodents. Animal Behaviour 94:27-34. https://doi.org/10.1016/j.anbehav.2014.05.008Links ]

B38 Sobrero, R., A. Ly Prieto, & L. A. Ebensperger. 2014b. Activity, overlap of range areas, and sharing of resting locations in the moon-toothed degu, Octodon lunatus. Journal of Mammalogy 95:91-98. https://doi.org/10.1644/13-mamm-a-144.1Links ]

B39 Swihart, R. K., & N. A. Slade. 1985. Testing for independence of observations in animal movements. Ecology 66:1176-1184. https://doi.org/10.2307/1939170Links ]

B40 Tang-Martínez, Z. 2003. Emerging themes and future challenges: Forgotten rodents, neglected questions. Journal of Mammalogy 84:1212-1227. https://doi.org/10.1644/ble-015Links ]

B41 Taraborelli, P. 2008. Vigilance and foraging behaviour in a social desert rodent, Microcavia australis (Rodentia Caviidae). Ethology Ecology & Evolution 20:245-256. https://doi.org/10.1080/08927014.2008.9522524Links ]

B42 Verzi, D. H., M. M. Díaz, & R. M. Barquez. 2015. Family Octodontidae Waterhouse, 1839. Mammals of South America, Rodents (J. L. Patton, U. F. J. Pardiñas & G. D’ Elía, eds.). The University of Chicago Press, Chicago and London. [ Links ]

B43 Whitehead, H. 2008. Analyzing animal societies: quantitative methods for vertebrate social analysis. University of Chicago Press, Chicago, Illinois. [ Links ]

B44 Whitehead, H. 2009. SOCPROG programs: analysing animal social structures. Behavioral Ecology and Sociobiology 63:765-778. https://doi.org/10.1007/s00265-008-0697-yLinks ]

Recibido: 27 de Julio de 2018; Aprobado: 14 de Diciembre de 2018