Mastozoología Neotropical ISSN: 0327-9383 ISSN: 1666-0536 [email protected] Sociedad Argentina para el Estudio de los Mamíferos Argentina

Frugone, María José; Correa, Loreto A; Sobrero, Raúl ACTIVITY AND GROUP-LIVING IN THE PORTER’S ROCK RATS, Aconaemys porteri Mastozoología Neotropical, vol. 26, núm. 2, 2019, Julio-, pp. 487-492 Sociedad Argentina para el Estudio de los Mamíferos Tucumán, Argentina

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

María José Frugone1, Loreto A. Correa2,3 and Raúl Sobrero4

1Instituto de Ecología y Biodiversidad, Departamento de Ciencias Ecológicas, Facultad de Ciencias, Universidad de Chile, Santiago, Chile. 2Escuela de Medicina Veterinaria, Facultad de Ciencias. Universidad Mayor, Santiago, Chile. 3Departamento de Ecología, Facultad de Ciencias, Ponticia Universidad Católica de Chile. 4Laboratorio de Ecología de Enfermedades, Instituto de Ciencias Veterinarias del Litoral (ICiVet-Litoral), Universidad Nacional del Litoral (UNL) / Consejo Nacional de Investigaciones Cientícas y Técnicas (CONICET), Esperanza, Argentina. [Correspondence: Raúl Sobrero ]

ABSTRACT. We provide the rst 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.

RESUMEN. Patrón de actividad y comportamiento social de Aconaemys porteri. 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.

Key words: activity patterns, group-living, home range, Octodontidae.

Palabras clave: ámbito de hogar, patrón de actividad, Octodontidae, vida en grupo.

Among caviomorph rodents, social organization Thus, caviomorphs oer unique opportunities to ranges from solitary living to social forms, in which examine factors associated with variation in social individuals interact frequently, overlap their range organization across and within species (e.g. Maher & areas, and share resting locations (e.g. Ebensperger Burger 2011). Within caviomorphs, the octodontids et al. 2004; Ebensperger & Hayes 2008). These ro- (Octodontidae) comprises 16 extant species grouped dents include species that dier morphologically and in six living genera with surface-dwelling (Octomys), physiologically and use a great diversity of habitats fossorial (Octodontomys, Octodon, Tympanoctomys), (Verzi et al. 2015) which seems to covary with group- completely subterranean (Spalacopus) and semisub- living (Lacey & Ebensperger 2007). terranean (Aconaemys) habits (Ojeda et al. 1996; Gallardo et al. 2007; Lessa et al. 2008).

Recibido 27 de julio 2018. Aceptado 14 diciembre 2018. Editor asociado: I. Tomasco 488 Mastozoología Neotropical, 26(2):487-492 Mendoza, 2019 M. J. Frugone et al. hp://www.sarem.org.ar – hp://www.sbmz.org

Aconaemys or rock rats used open areas and the Fundo San Pablo de Tregua (39°35’S, 72°05’W), forests, on both eastern and western slopes of the Panguipulli, Chile. Andes (Pearson 1984; Gallardo & Mondaca 2002). San Pablo de Tregua is characterized by a dry and Three species are currently recognized: A. fus- short summer where mean annual precipitation is cus, A. sagei, and A. porteri (Gallardo & Mondaca around 5000 mm, and ambient temperature ranges 2002). Aconaemys fuscus inhabits highland forests from 5°C to 20°C (Alvarez & Lara 2008; Guevara of Araucaria araucana (Araucariaceae) and sandy et al. 2015). The site consisted of Valdivian rain- areas (Muñoz-Pedreros 2000), while A. sagei inhabits forest where dominant tree species were coigüe ungrazed bunchgrass and Nothofagus forest with (Nothofagus dombeyi), raulí (N. alpina), and tepa bamboo cover (Pearson 1984). Aconaemys porteri (Laureliopsis philippiana) (Schlegel & Donoso 2008). is distributed from Volcán Villarrica to Puyehue Animals were captured using a combination of 172 in Chile, and from both Parque Nacional Lanín (14 x 14 x 40-cm) Tomahawk (model 201; Tomahawk and Parque Nacional Nahuel Huapi in Argentina Live Trap Company, Hazelhurst, Wisconsin) and (Pearson 1984; 1995; Gallardo & Reise 1992; Gallardo leg-hold traps during 8 days in January, 2011. & Mondaca 2002). In these areas, Porter’s rock Qualitatively, putative nest sites used by A. porteri rats inhabit dense bamboo and southern beech rain- are structurally similar to burrows and resting lo- forests (Pearson 1983; 1984; Gallardo & Reise 1992). cations of O. degus (Fulk 1976; Lessa et al. 2008), consisting mainly of oblique tunnels connecting the Evaluating the extent of the social behavior of surface to the nest, with several branches and active Aconaemys remains critical to determine whether openings (4-5) associated to patches of perennial ecological conditions and species-specic attributes bamboo (Chusquea quila), covering part of the bur- drove the evolution of group living in the octodon- row system. We dened rats burrow as areas of tids. A recent comparative analysis suggested that 1.5 m2 covered by vegetation, in which we found social behavior in caviomorphs was gained and lost signs of A. porteri presence (i.e., feces) and where repeatedly, perhaps originating from an ancestral radiocollared individuals were found during trap- species that was socially exible, and where the ping and telemetry periods. Ground mounds were loss of group-living has been associated to the use sometimes found outside openings and fresh feces of habitats with high plant cover (Sobrero et al. in burrows openings allow us to determine if each 2014a). Moreover, robust knowledge about multiple burrow system was active. ecological factors (distribution of resources such as Moreover, in two occasions rats’ vocalizations food, predation risk, and soil conditions associated were listened (but not recorded and analyzed) during with digging burrows or nesting sites) surely will trapping. We placed traps near burrow openings contribute to build a theory of sociobiology that and inside patches with high bamboo cover and is closer and more consistent with the diversity of baited them with rolled oats, cereals and sunower mammalian social behavior (Tang-Martínez 2003; seeds. Traps were opened 08:00h and closed 00:00h Hayes et al. 2007). Taken together, establishing and checked every hour. During each capture, we how group-living varies across species is critical recorded sex, body mass and reproductive status, and for comparative studies to examine the origin and each animal was marked with an ear tag (Monel 1005- the adaptive value of this behavior (Blumstein & 1; National Band and Tag Co., Newport, Ky., USA). Armitage 1998; Ebensperger 2001; Ebensperger & Low number of rats represents the species at a nor- Blumstein 2006). The scarce available and anecdotal mal density, based on knowledge about population evidence suggests that Porter’s rock rats lives in ecology of other caviomorph rodents (e.g. Cassini small groups in communal burrow systems and both 1991; Ebensperger et al. 2008). All adult-sized individ- diurnal and nocturnal activity has been recorded uals (N = 5) were tted with a radiocollar weighing (Pearson 1983; Verzi et al. 2015). Also, in captivity, in- 7–9 g (BR radiocollars; AVM Instrument Co., Colfax, dividuals displayed a high tolerance for conspecics California) with unique pulse frequencies. Weight of (Verzi et al. 2015); however it has not been tested radio-collars represented about 4–5% of body adult in natural populations and laboratory conditions weight (e.g. Ebensperger et al. 2008). At the end of can modify behavior of captive individuals (Calisi & our study all radiocollared animals were recaptured Bentley 2009). and radiocollars were removed (Ebensperger et al. Our aim in this research note is to report data 2004; 2012). of the activity patterns and social behavior of free- During 5 days and nights it was performed hom- living A. porteri. The study site was located at ing technique every two hours, to determine resting BEHAVIOR AND ECOLOGY OF Aconaemys porteri (OCTODONTIDAE) 489

locations or putative nest places, using LA 12-Q The range area was determined from locations receiver (for radiocollars tuned to 150.000–151.999 recorded through triangulation and included animal MHz frequency; AVM Instrument Co., Colfax, resting locations. While triangulation is thought California) and a handheld 3-element yagi antenna to interfere less with the activity of radio-collared (AVM instrument Co., Colfax, California). Once animals compared with homing (Kenward 2001; located, the position of each animal was marked Ebensperger & Blumstein 2006), the topography and with agging material coded for individuals. Each cover type of our study site precluded the use of long- radiox location was then referenced twice with a range radio-xings because of signal bounce. Thus, Garmin portable global positioning system (Garmin although we used the homing technique, we previ- International Inc., Olathe, Kansas), precision always ously trained ourselves to locate animals quickly to was within 5 m. The determination of group com- minimize disrupting their navigation or locomotion position required the compilation of a symmetric behavior (see Ebensperger et al. 2008). Data points similarity matrix of pairwise association of the rest- from each individual were mapped using the 95% ing locations of all adult animals during homing minimum convex polygon algorithm in Ranges 6 (Whitehead 2008). Social organization was quanti- (Kenward et al. 2003). Pairwise estimates of the per- ed based on the number and sex composition of cent overlap between polygons for dierent animals adult members in a social group (Ebensperger & also were calculated using Ranges 6. We compared Hayes 2016). Thus, we conducted a hierarchical clus- the mean size (in m2) of range areas and percent ter analysis of the association matrix in SOCPROG range overlap by male and female Porter’s rock rats software (Whitehead 2009). with Mann–Whitney U-tests. We used Wilcoxon matched-pair tests to compare percent overlap in range areas of individuals assigned to the same To estimate daily activity patterns and range areas, burrow location associations and percent overlap we recorded locations hourly of all radiocollared that these individuals had with individuals assigned animals for 5 days and 4 nights in 2011, at nighttime to dierent associations in January 2011. All sta- (21:00-07:00 h) and daytime (07:00-21:00 h). Sunrise tistical analyses were calculated using Statistica 7.0 occurred at approximately 06:30 h, whereas sunset (StatSoft Inc. 1984–2004) and results are reported as occurred at 20:30 h. The spatial location of animals mean ± SE. was determined using triangulation (Kenward 2001). We used 2 LA 12-Q receivers, each connected to a Animals used 2 to 9 burrow locations, namely null peak antenna system (AVM Instrument Co.). where one or more rats were found repeatedly during Every null peak system had four 7-element yagi total radioscans (n=22). We recorded 21 scans in antennas. Distance between antenna stations was which animals shared burrow or resting locations. about 85 m. Bearings from both antenna stations Of these, 20 observations involved male–female were then transformed into x–y locations with the pairs, and 1 involved 2 males–1 female associations. software Locate II (Nams 1990). We calculated the Distance moved between radioscans was variable distance traveled (in m), between successive scans through time of day or night (Fig. 1). The distance as a measure of aboveground activity of the Porter’s moved between any two consecutive radio xes rock rats. The same individuals were monitored during the night averaged 49 ± 28 m (n = 5, range: throughout consecutive days and nights. As a re- 1-249 m), and was 1.4 times larger than distance sult, locations recorded at 24-h intervals were not moved during day (36 ± 25 m, range: 0.5-425 m). independent of one another. Consequently, and However, this dierence was not statistically sig- for statistical analyses, we divided the entire data nicant (F8,9 = 0.420, P = 0.535). Five individuals collection period into 5 days and 4-night cycles, were radio-tracked during the day and night, which dened on the basis of sunrise and sunset at study provided an average of 98 useful radio xes per ani- site. For the daytime portion of each activity cycle, mal. Spatial overlap among radio-collared Porter’s we calculated the mean distance travelled for each rock rats was relatively low (20 ± 5%) ranging from radiocollared individual. We used a similar approach 3% to 34% (Fig. 2). If data from all individuals for the nighttime portion of the activity cycle. As are combined, the size of range areas averaged a result, each radiocollared individual contributed 48 ± 13 m2 (n = 5). When sex was examined, females 2 dependent data points to our analysis of activity. tended to range over larger areas (51 ± 3 m2, n = 3) We used repeated-measures analysis of variance to than males (45 ± 39 m2, n = 2), a non-statistically examine the eect of activity time (day versus night) signicant dierence (Mann–Whitney U test, z = on individual activity of males and females. 3.0, p > 0.10). 490 Mastozoología Neotropical, 26(2):487-492 Mendoza, 2019 M. J. Frugone et al. hp://www.sarem.org.ar – hp://www.sbmz.org

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 ve Porter’s rock rats (Aconaemys porteri) during the activity period. The arrow signals the geographic north. BEHAVIOR AND ECOLOGY OF Aconaemys porteri (OCTODONTIDAE) 491

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