Raúl Sobrero

Sociality, glucocorticoids and direct fitness in the communally rearing rodent, Octodon degus

While ecological causes of sociality (or group living) have been identified, proximate mechanisms remain less clear. Recently, close connections between sociality, glucocorticoid hormones (cort) and fitness have been hypothesized. In particular, cort levels would reflect a balance between fitness benefits and costs of group living, and therefore baseline cort levels would vary with sociality in a way opposite to the covariation between sociality and fitness. However, since reproductive effort may become a major determinant of stress responses (i.e., the cort-adaptation hypothesis), cort levels might also be expected to vary with sociality in a way similar to the covariation between sociality and fitness. We tested these expectations during three years in a natural population of the communally rearing degu, Octodon degus. During each year we quantified group membership, measured fecal cortisol metabolites (a proxy of baseline cort levels under natural conditions), and estimated direct fitness. We recorded that direct fitness decreases with group size in these animals. Secondly, neither group size nor the number of females (two proxies of sociality) influenced mean (or coefficient of variation, CV) baseline cortisol levels of adult females. In contrast, cortisol increased with per capita number of offspring produced and offspring surviving to breeding age during two out of three years examined. Together, our results imply that variation in glucocorticoid hormones is more linked to reproductive challenge than to the costs of group living. Most generally, our study provided independent support to the cort-adaptation hypothesis, according to which reproductive effort is a major determinant, yet temporally variable, influence on cort-fitness covariation.

Burrow limitations and group living in the communally rearing rodent, Octodon degus

Abstract Group living is thought to evolve whenever individuals attain a net fitness advantage due to reduced predation risk or enhanced foraging efficiency, but also when individuals are forced to remain in groups, which often occurs during high-density conditions due to limitations of critical resources for independent breeding. The influence of ecological limitations on sociality has been studied little in species in which reproduction is more evenly shared among group members. Previous studies in the caviomorph rodent Octodon degus (a New World hystricognath) revealed no evidence that group living confers an advantage and suggest that burrow limitations influence formation of social groups. Our objective was to examine the relevance of ecological limitations on sociality in these rodents. Our 4-year study revealed no association between degu density and use of burrow systems. The frequency with which burrow systems were used by degus was not related to the quality of these structures; only in 1 of the 4 years did the frequency of burrow use decrease with decreasing abundance of food. Neither the number of females per group nor total group size (related measures of degu sociality) changed with yearly density of degus. Although the number of males within social groups was lower in 2008, this variation was not related clearly to varying density. The percentage of females in social groups that bred was close to 99% and did not change across years of varying density. Our results suggest that sociality in degus is not the consequence of burrow limitations during breeding. Whether habitat limitations contribute to variation in vertebrate social systems is discussed.

Towards an integrative model of sociality in caviomorph rodents

Abstract In the late 1990s and early 2000s it was recognized that behavioral ecologists needed to study the sociality of caviomorph rodents (New World hystricognaths) before generalizations about rodent sociality could be made. Researchers identified specific problems facing individuals interested in caviomorph sociality, including a lack of information on the proximate mechanisms of sociality, role of social environment in development, and geographical or intraspecific variation in social systems. Since then researchers have described the social systems of many previously understudied species, including some with broad geographical ranges. Researchers have done a good job of determining the role of social environments in development and identifying the costs and benefits of social living. However, relatively little is known about the proximate mechanisms of social behavior and fitness consequences, limiting progress toward the development of integrative (evolutionary-mechanistic) models for sociality. To develop integrative models behavioral ecologists studying caviomorph rodents must generate information on the fitness consequences of different types of social organization, brain mechanisms, and endocrine substrates of sociality. We review our current understanding and future directions for research in these conceptual areas. A greater understanding of disease ecology, particularly in species carrying Old World parasites, is needed before we can identify potential links between social phenotypes, mechanism, and fitness.

Phylogeny modulates the effects of ecological conditions on group living across hystricognath rodents

The evolution of group living in animals has been linked both to ecological and phylogenetic drivers or constraints. However, available evidence supporting each factor remains equivocal. In this study, we evaluated the influence of both scenarios across 98 species of hystricognath rodents using phylogenetic approaches that included a statistical model of trait evolution. We first estimated the hypothetical ancestral forms of several clades within the hystricognath phylogeny. Secondly, we examined evolutionary correlations between habitat conditions in terms of vegetation cover and group living to determine general macroevolutionary trends on social behaviour. Given the contrasting modes of life that characterize these rodents, we also examined whether subterranean habits influence group living. Our results support the hypothesis that group living in Old World Phiomorpha and New World Caviomorpha had an early origin. The best-fitted and well-supported model of evolution for group living was the equal rate model, implying an equal rate of gains and losses of group living. Logistic regression analyses that included phylogenetic relationships revealed significant correlation between habitats used (i.e. vegetation cover) and absence or presence of group living. The loss of group living across the phylogeny was associated with the use of closed habitat conditions. No association was found between changes in group living and subterranean mode of life. We conclude that ancestor–descendant relationships mediate the effect of changes in habitat use on the evolution of group living across -hystricognath rodents.

Relatedness does not predict vigilance in a population of the social rodent Octodon degus

The possibility that social foragers adjust and coordinate their scanning activity when in the presence of close relatives to attain inclusive fitness benefits remains controversial and scarcely examined. To this aim, we first tested the null hypothesis of no association between foraging individuals of the diurnal rodent, Octodon degus and their pairwise relatedness (six microsatellite loci), under natural conditions. Secondly, we examined the influence of relatedness on scan effort (percent overlapping) and temporal distribution of scanning using linear regression. Finally, we evaluated whether temporal distributions of scanning were significantly lower (coordination) or higher (synchrony) than random expectations using bootstrapping. We found that pairwise relatedness between focal degus and their foraging partner did not influence the scan effort or the temporal distribution of scanning. These original, field-based findings imply that vigilance behavior in socially foraging degus is unlikely to be kin-selected and adds to results from previous lab studies in that kinship remains a poor predictor of social behavior in these animals. Overall, our study adds to others revealing that kin selection may not have had an impact on aspects of social behavior such as vigilance during social foraging.

Octomys mimax (Rodentia: Octodontidae)

Abstract Octomys mimax O. Thomas, 1920, the viscacha rat, is a surface-dwelling rodent, endemic to the Monte biome of western Argentina. The genus Octomys is monotypic and represents a basal clade within the family Octodontidae. O. mimax is strictly herbivorous and is found in low-elevation arid environments characterized by galleries or rock formations. In these environments, O. mimax nests within rock crevices located inside ravines with relatively low vegetation cover. O. mimax is solitary and mostly nocturnal. Nocturnal activity coupled with a basal metabolic rate and thermal conductance lower than expected for body mass of O. mimax probably represent strategies for water conservation in this species.

Patterns of variation in the tympanic bulla of tuco-tucos (Rodentia, Ctenomyidae, Ctenomys)

As subterranean rodents live in burrows and are constrained by the physics of their environment to vocalize, mainly in low frequencies, their expanded middle ear cavities are associated with enhanced lower-frequency hearing. Previous literature has widely acknowledged inflated tympanic bulla as a character to be found in the majority of the Ctenomys species. To explore the morphology of Ctenomys tympanic bulla, we studied a sample of 669 skulls, obtained from 21 species, for tympanic bulla size, volume, and internal structure. The study determined that bullar inflation does not seem to be the rule in Ctenomys and that the relationship between bullar size (volume) and skull size do not correspond to the phylogeny based on cytochrome b sequences thus probably being a species-specific adaptive characteristic. We also found that the internal bullar structure differs between taxa, depending on the relative contributions of cancellous (alveolar) and septate patterns to the partitioning of the bulla.

Expensive brains: "brainy" rodents have higher metabolic rate

Brains are the centers of the nervous system of animals, controlling the organ systems of the body and coordinating responses to changes in the ecological and social environment. The evolution of traits that correlate with cognitive ability, such as relative brain size is thus of broad interest. Brain mass relative to body mass (BM) varies among mammals, and diverse factors have been proposed to explain this variation. A recent study provided evidence that energetics play an important role in brain evolution (Isler and van Schaik, 2006). Using composite phylogenies and data drawn from multiple sources, these authors showed that basal metabolic rate (BMR) correlates with brain mass across mammals. However, no such relationship was found within rodents. Here we re-examined the relationship between BMR and brain mass within Rodentia using a novel species-level phylogeny. Our results are sensitive to parameter evaluation; in particular how species mass is estimated. We detect no pattern when applying an approach used by previous studies, where each species BM is represented by two different numbers, one being the individual that happened to be used for BMR estimates of that species. However, this approach may compromise the analysis. When using a single value of BM for each species, whether representing a single individual, or available species mean, our findings provide evidence that brain mass (independent of BM) and BMR are correlated. These findings are thus consistent with the hypothesis that large brains evolve when the payoff for increased brain mass is greater than the energetic cost they incur.

Activity, overlap of range areas, and sharing of resting locations in the moon-toothed degu, Octodon lunatus

Abstract The evolution of sociality across octodontid rodents remains puzzling. Although basal species are solitary living, the most derived octodontids studied so far are social, implying that sociality evolved recently from solitary-living ancestors. However, the social behavior of some octodontids remains anecdotal. We aimed to provide the 1st systematic data on activity, space use, and social behavior of the moon-toothed degu (Octodon lunatus), a derived octodontid rodent. We used livetrapping and radiotelemetry to monitor patterns of aboveground activity, aboveground range areas and overlap, and use of resting locations in a coastal population in north-central Chile. Activity of O. lunatus was statistically similar during nighttime and daytime, implying no clear diurnal or nocturnal activity. During daytime the animals used resting locations that were associated with high shrub cover and Pouteria splendens. Radiocollared males and females shared resting locations on multiple occasions. There was a nonsignificant trend in degus that used same resting locations to exhibit greater range overlap than degus using different resting locations. Associations based on resting locations revealed a total of 5 social groups. Taken together, these results indicate that adult O. lunatus exhibit some sociality, a finding consistent with a trend in which group living is more frequent in the most derived compared with basal octodontids. Resumen Nuestro conocimiento sobre la evolución del comportamiento social en roedores octodóntidos es aún fragmentario. La información disponible indica que las especies filogenéticamente basales son solitarias, mientras que las más derivadas tienden a ser sociales. Sin embargo, la información sobre la estructura social disponible para varias especies es anecdótica, lo cual dificulta el establecimiento de conclusiones robustas sobre la evolución del comportamiento social en este clado. Este es el primer estudio que cuantifica la actividad, uso del espacio, y comportamiento social del degú costino (Octodon lunatus), una especie derivada de octodóntido. Durante noviembre y diciembre de 2010 y 2011 se utilizaron métodos de captura–recaptura y telemetría para cuantificar el patrón diario de actividad superficial, ámbitos de hogar, solapamientos entre ámbitos de hogar, y uso compartido de parches de descanso y nidificación en una población costera localizada en el centro-norte de Chile. La actividad de O. lunatus, medida como desplazamientos individuales entre localizaciones consecutivas, mostró una tendencia estadísticamente no significativa a ser mayor en horas de la noche. Durante el día los animales usaron 1 a 3 sitios de descanso y anidamiento asociados con una alta cobertura arbustiva, donde Pouteria splendens (lúcumo) fue la especie dominante. Machos y hembras compartieron estos sitios de descanso en múltiples ocasiones. El solapamiento entre los ámbitos de hogar tendió a ser mayor en animales que además compartieron sitios de descanso comparado con animales que no compartieron estos sitios. En base al uso compartido de refugios se identificó 1 grupo social en 2010 y 4 grupos en 2011. La composición de estos grupos fue de 1 a 3 hembras adultas y de 1 a 2 machos adultos (2 a 4 adultos en total). Globalmente, los resultados indicaron que O. lunatus muestra algún grado de sociabilidad, observación que apoya una tendencia en la cual el comportamiento social es más frecuente en especies filogenéticamente derivadas de octodóntidos.

Effects of Habitat and Social Complexity on Brain Size, Brain Asymmetry and Dentate Gyrus Morphology in Two Octodontid Rodents

Navigational and social challenges due to habitat conditions and sociality are known to influence dentate gyrus (DG) morphology, yet the relative importance of these factors remains unclear. Thus, we studied three natural populations of O. lunatus (Los Molles) and Octodon degus (El Salitre and Rinconada), two caviomorph species that differ in the extent of sociality and with contrasting vegetation cover of habitat used. The brains and DG of male and female breeding degus with simultaneous information on their physical and social environments were examined. The extent of sociality was quantified from total group size and range area overlap. O. degus at El Salitre was more social than at Rinconada and than O. lunatus from Los Molles. The use of transects to quantify cover of vegetation (and other physical objects in the habitat) and measures of the spatial behavior of animals indicated animal navigation based on unique cues or global landmarks is more cognitively challenging to O. lunatus. During lactation, female O. lunatus had larger brains than males. Relative DG volume was similar across sexes and populations. The right hemisphere of male and female O. lunatus had more cells than the left hemisphere, with DG directional asymmetry not found in O. degus. Degu population differences in brain size and DG cell number seemed more responsive to differences in habitat than to differences in sociality. Yet, large-sized O. degus (but not O. lunatus) that ranged over larger areas and were members of larger social groups had more DG cells per hemisphere. Thus, within-population variation in DG cell number by hemisphere was consistent with a joint influence of habitat and sociality in O. degus at El Salitre.