Bite hard: Linking cranial loading mechanics to ecological differences in gnawing behavior in caviomorph rodents.

The mammalian skull is very malleable and has notably radiated into highly diverse morphologies, fulfilling a broad range of functional needs. Although gnawing is relatively common in mammals, this behavior and its associated morphology are diagnostic features for rodents. These animals possess a very versatile and highly mechanically advantageous masticatory apparatus, which, for instance, allowed caviomorph rodents to colonize South America during the Mid-Eocene and successfully radiate in over 200 extant species throughout most continental niches. Previous work has shown that differences in bite force within caviomorphs could be better explained by changes in muscle development than in mechanical advantages (i.e., in cranial overall morphology). Considering the strong bites they apply, it is interesting to assess how the reaction forces upon the incisors (compression) and the powerful adductor musculature pulling (tension) mechanically affect the cranium, especially between species with different ecologies (e.g., chisel-tooth digging). Thus, we ran finite element analyses upon crania of the subterranean Talas' tuco-tuco Ctenomys talarum, the semi-fossorial common degu Octodon degus, and the saxicolous long-tailed chinchilla Chinchilla lanigera to simulate: (A) in vivo biting in all species, and (B) rescaled muscle forces in non-ctenomyid rodents to match those of the tuco-tuco. Results show that the stress patterns correlate with the mechanical demands of distinctive ecologies, on in vivo-based simulations, with the subterranean tuco-tuco being the most stressed species. In contrast, when standardizing all three species (rescaled models), non-ctenomyid models exhibited a several-fold increase in stress, in both magnitude and affected areas. Detailed observations evidenced that this increase in stress was higher in lateral sections of the snout and, mainly, the zygomatic arch; between approximately 2.5-3.5 times in the common degu and 4.0-5.0 times in the long-tailed chinchilla. Yet, neither species, module, nor simulation condition presented load factor levels that would imply structural failure by strong, incidental biting. Our results let us conclude that caviomorphs have a high baseline for mechanical strength of the cranium because of the inheritance of a very robust "rodent" model, while interspecific differences are associated with particular masticatory habits and the concomitant level of development of the adductor musculature. Especially, the masseteric and zygomaticomandibular muscles contribute to >80% of the bite force, and therefore, their contraction is responsible for the highest strains upon their origin sites, that is, the zygomatic arch and the snout. Thus, the robust crania of the subterranean and highly aggressive tuco-tucos allow them to withstand much stronger forces than degus or chinchillas, such as the ones produced by their hypertrophied jaw adductor muscles or imparted by the soil reaction.

Ontogenetic skull variation in a shovel-headed amphisbaenian species.

Leposternon microcephalum is a species belonging to the Amphisbaenia, a group of burrowing reptiles. Amphisbaenia present various morphological and physiological adaptations that allow them to penetrate the ground and live underground, through a system of galleries and permanent chambers that they build themselves. Among the morphological adaptations in this group, those of the skull stand out as it serves as the main excavation tool. Four basic skull shapes are recognized: rounded, keeled, shovel-shaped, and spade-shaped. The skull of L. microcephalum belongs to this last type, which is considered the most specialized. The species inhabits soils that are highly compacted and difficult to penetrate. Among the species of Leposternon present in South America, L. microcephalum has the widest distribution, being found in all Brazilian biomes and neighboring countries such as Bolivia, Argentina, Paraguay, and Uruguay. The analysis of the skull of this species was carried out using three-dimensional geometric morphometrics (3D-GMM), a technique that allows comparative analysis, through robust statistical methods, of shape and its variations, using Cartesian coordinate data from a configuration of homologous landmarks. The technique allows the size and shape components of a structure to be analyzed separately. From an ontogenetic point of view, this methodology had also been used to investigate variations in Cynisca leucura, a member of the Amphisbaenidae with a rounded head. Our hypothesis is that the patterns of morphological differentiation in the skull, mainly in the intermediate and occipital regions, are similar in different Amphisbaenia species. Therefore, the objective of this study was to analyze cranial morphological variations in an ontogenetic series of L. microcephalum using 3D-GMM. Computed Tomographic scans of 13 specimens were analyzed: juveniles (N = 8) and adults (N = 5), based on 20 landmarks that characterize the skull. Principal components and regression analyses between shape (dependent variable) and size (independent variable) showed a clear difference between the cranial morphological pattern of juvenile individuals and that of adults. For instance, young specimens tend to have a dorsoventrally tall neurocranium, with the tip of the snout more anteriorly oriented and its dorsal border subtly curved. Dorsally, the parietal region is thicker and smoothly dome-shaped in juveniles. As in C. leucura, the variation was strongly correlated with the size change from juvenile to adult, indicating a dominant role for ontogenetic allometry in determining skull shape.

Shape analysis of the preorbital bar in caviomorph rodents.

The highly specialised masticatory apparatus of rodents raises interesting questions about how their skull withstands the intensive and sustained forces produced by biting on hard items. In these mammals, major systematics were explored for a long time based on the adductor muscles' architecture and the related bony structures. The infraorbital foramen stands out, where a hypertrophied head of the zygomaticomandibular muscle passes through-in hystricomorphous rodents-as a direct consequence of the lateral and posterior shift of the preorbital bar. Interestingly, this bar moved laterally and backwards-enlarging the foramen-but it never disappeared throughout evolution, even showing morphological convergence among rodents. Previous research proposed this bar as behaving mechanically similar to the postorbital bar in ungulates, i.e., a safety structure against torsion stress while chewing. We analysed its morphology by mathematically modelling it under bending and torsion scenarios (linearly and elliptically shaped, respectively), and as for biting load propagation (catenary curve). Although the preorbital bar primarily seems to be shaped for withstanding torsional stress (as the postorbital bar in ungulates) and as an escaping point for force propagation, these forces are not a consequence of chewing and grinding foods, but preventing the zygomatic arch from failing when the powerful laterally-displaced jaw adductor muscles are pulling the dentary upwards at biting.

Functional morphology and fiber types of the masseter muscles of two caviomorph rodents with contrasting lifestyles, Ctenomys talarum (Ctenomyidae) and Cavia aperea (Caviidae).

The aim of this work is the analysis of histochemical and morphometric properties of the masseter muscles of Ctenomys talarum and Cavia aperea. The former belongs to a subterranean rodent clade, Ctenomyidae, which has evolved a robust masticatory apparatus adapted to chisel-tooth digging and processing of abrasive grasses; C. aperea belongs to the family Caviidae, with relatively graceful jaws and mandibular musculatures, consistent with less mechanically challenging diets. Adult males were captured, immediately transported to the laboratory, and euthanized in a CO2 chamber. The musculus masseter superficialis and musculus masseter profundus on the left side of the animals were used to analyze the histochemical composition of the fiber types treated with myosin adenosine triphosphatase, succinate dehydrogenase and periodic acid Schiff. The mean fiber diameters, relative areas, and frequencies of each muscle fiber type were calculated. The mm. masseter superficialis and masseter profundus on the right side were used to measure the physiological cross-sectional area (PCSA). Based on this measurement, the internal force (F) was estimated. In the m. masseter profundus of both species and in the m. masseter superficialis of C. aperea intermediate fast oxidative-glycolytic fibers (FOGi) predominated. In the mm. masseter superficialis and masseter profundus of C. talarum the relative area of fast glycolytic (FG) fibers was greater than that of the muscles of C. aperea, whose main muscle fiber component is FOGi fibers. When corrected for body mass differences, PCSA was higher for the mm. masseter superficialis of C. talarum. This and the larger relative area of FG fibers, probably contributes to the exertion of large bite forces in C. talarum, as measured in previous studies.

Functional morphology and identity of the thenar pad in the subterranean genus Ctenomys (Rodentia, Caviomorpha).

As in many other fossorial tetrapods, the most obvious adaptations to scratch-digging in the subterranean tuco-tuco (Rodentia, Ctenomyidae, Ctenomys) are found in the hands, which among other adaptations, present the mesaxonic condition; i.e. the central digits are more developed, and also their claws, which are curved and elongated. The thumb is atrophied and aligned with the rest of the digits, showing a small and flat claw. This configuration of digits and claws seems to be in accordance with what it is expected for rodents: rudimentary movements when handling food items. However, on the palmar side of the hand, tuco-tucos have several pads, the thenar (located under the thumb) being the most developed. In this study, we investigated the functional morphology of the thenar pad through different approaches: musculoskeletal anatomy, histology and functionality. The analysis of radiographs and clarified and double-stained hand samples of Ctenomys talarum and C. australis showed that the thenar pad is supported by a paddle-shaped bone that articulates with a protrusion in the scapholunate bone. This bone, flat and long, continues in a flat cartilaginous structure, with a shape similar to a claw. Dissections showed that the thenar pad has several associated muscles: the m. palmaris longus, the m. abductor pollicis longus, and a massive muscular complex located between the thumb and the thenar pad. By topology it might be inferred that this complex is formed by the m. abductor pollicis brevis, the m. flexor pollicis brevis and the m. adductor pollicis brevis. Longitudinal histological sections of the thenar pad stained with hematoxylin-eosin showed a thick layer of keratin at the distal end, external face. The observation of live specimens of C. talarum foraging on two food items of different size and filmed at 300 fps showed that the thenar pad acts as an opposable thumb, with digit-like movements. Tuco-tucos are able to perform more precise movements than expected, and to grasp and manipulate the food with one hand. In previous studies, it was suggested that the thenar pad was supported by a 'palmar ossicle', or 'prepollex' (= radial sesamoid bone). Our results suggest that this sesamoid underwent a radical change on its morphology, making the thenar pad a part of the food handling system in Ctenomys, so the thenar pad might be considered a 'false thumb', rather than a palmar pad. It is suggested to advance on the description and functional analysis of the thenar pad, redefining the structure, since the terms used so far to define it would not be accurate.

Mandible strength and geometry in relation to bite force: a study in three caviomorph rodents.

The monophyletic group Caviomorpha constitutes the most diverse rodent clade in terms of locomotion, ecology and diet. Caviomorph species show considerable variation in cranio-mandibular morphology that has been linked to the differences in toughness of dietary items and other behaviors, such as chisel-tooth digging. This work assesses the structural strength of the mandible of three caviomorph species that show remarkable differences in ecology, behavior and bite force: Chinchilla lanigera (a surface-dwelling species), Octodon degus (a semi-fossorial species) and Ctenomys talarum (a subterranean species). Finite element (FE) models of the mandibles are used to predict the stresses they withstand during incisor biting; the results are related to in vivo bite forces and interspecific variations in the mandibular geometries. The study concludes that the mandible of C. talarum is better able to withstand strong incisor bites. Its powerful adducting musculature is consistent with the notorious lateral expansion of the angular process and the masseteric crest, and the enhanced cortical bone thickness. Although it has a relatively low bite force, the mandible of O. degus also shows a good performance for mid-to-strong incisor biting, in contrast to that of C. lanigera, which exhibits, from a mechanical point of view, the worst performance. The mandibles of C. talarum and O. degus appear to be better suited to withstand stronger reaction forces from incisor biting, which is consistent with their closer phylogenetic affinity and shared digging behaviors. The contrast between the low in vivo bite force of C. lanigera and the relatively high estimations that result from the models suggests that its adductor musculature could play significant roles in functions other than incisor biting.

Cranial suture complexity in caviomorph rodents (Rodentia; Ctenohystrica).

Due to their flexibility, sutures are regions that experience greater strains than the surrounding rigid cranial bones. Cranial sutures differ in their degree of interdigitation or complexity. There is evidence indicating that a more convoluted suture better enables the absorption of high stresses coming from dynamic masticatory forces, and other functions. The Order Rodentia is an interesting clade to study this because of its taxa with diverse chewing modes. Due to repeated loading resulting from gnawing and grinding, energy absorption by the sutures might be a crucial factor in these mammals. Species within the infraorder Caviomorpha were chosen as a case study because of their ecomorphological and dietary diversity. This study compared five sutures from the rostrum and cranial vault across seven caviomorph families, and assessed their complexity by means of the relative length and fractal dimension. Across these rodents, cranial sutures are morphologically quite diverse. We found that the sutures connecting the rostrum with the vault were relatively more interdigitated than those in the cranial vault itself, especially premaxillofrontal sutures. Suture interdigitation was higher in species that display chisel-tooth digging and burrowing behaviors, especially in the families Ctenomyidae and Octodontidae, than those in families Dasyproctidae and Cuniculidae, which have more gracile masticatory systems. The reconstruction of the ancestral character state, on family and species phylogeny, points toward low suture interdigitation (i.e., low length ratio) as a likely ancestral state for interfrontal, premaxillofrontal and maxillofrontal sutures. Interspecific differences in suture morphology shown here might represent adaptations to different mechanical demands (i.e., soft vs. tough foods) or behaviors (e.g., chisel-tooth digging), which evolved in close association with the diverse environments occupied by caviomorph rodents.

Analysis of arch-like bones: The rodent mandible as a case study.

Bone strength is determined by the mechanical properties of bone material, and the size and shape of the whole bone, i.e., its architecture. The mandible of vertebrates has been traditionally regarded as a beam oriented in relation to main masticatory loads, i.e., the longer dimension of its cross-section being parallel to the load. Rodents follow this pattern but, in addition, their mandible possesses an intriguing arch-like shape that is apparent when seen in the lateral view. Little attention was given to the structural capacity of this trait. The advantage of an arch is that it can withstand a greater load than a horizontal beam. The objective of this study was to model the rodent mandible like an arch to evaluate its structural strength. The bending moment in an arch-like mandible was 15-25% lower with respect to a beam-like mandible. Further, bending varies with mandible "slenderness" and incisor procumbency, a functionally relevant rodent trait. In the rodent Ctenomys talarum (Caviomorpha; Ctenomyidae), bone stress was substantially reduced when the mandible was modeled as an arch-like structure as compared with a beam-like structure, and safety factors were 15-34% higher. The shape of rodents' mandible might confer a functional advantage to high and repeatedly applied loads resulting from a unique feeding mode: gnawing. J. Morphol. 277:879-887, 2016. © 2016 Wiley Periodicals, Inc.

Postnatal ontogeny of limb proportions and functional indices in the subterranean rodent Ctenomys talarum (Rodentia: Ctenomyidae).

Burrow construction in the subterranean Ctenomys talarum (Rodentia: Ctenomyidae) primarily occurs by scratch-digging. In this study, we compared the limbs of an ontogenetic series of C. talarum to identify variation in bony elements related to fossorial habits using a morphometrical and biomechanical approach. Diameters and functional lengths of long bones were measured and 10 functional indices were constructed. We found that limb proportions of C. talarum undergo significant changes throughout postnatal ontogeny, and no significant differences between sexes were observed. Five of six forelimb indices and two of four hindlimb indices showed differences between ages. According to discriminant analysis, the indices that contributed most to discrimination among age groups were robustness of the humerus and ulna, relative epicondylar width, crural and brachial indices, and index of fossorial ability (IFA). Particularly, pups could be differentiated from juveniles and adults by more robust humeri and ulnae, wider epicondyles, longer middle limb elements, and a proportionally shorter olecranon. Greater robustness indicated a possible compensation for lower bone stiffness while wider epicondyles may be associated to improved effective forces in those muscles that originate onto them, compensating the lower muscular development. The gradual increase in the IFA suggested a gradual enhancement in the scratch-digging performance due to an improvement in the mechanical advantage of forearm extensors. Middle limb indices were higher in pups than in juveniles-adults, reflecting relatively more gracile limbs in their middle segments, which is in accordance with their incipient fossorial ability. In sum, our results show that in C. talarum some scratch-digging adaptations are already present during early postnatal ontogeny, which suggests that they are prenatally shaped, and other traits develop progressively. The role of early digging behavior as a factor influencing on morphology development is discussed.

Another one bites the dust: bite force and ecology in three caviomorph rodents (Rodentia, Hystricognathi).

Mammals have developed sophisticated strategies adapting to particular locomotor modes, feeding habits, and social interactions. Many rodent species have acquired a fossorial, semi-fossorial, or even subterranean life-style, converging on morphological, anatomical, and ecological features but diverging in the final arrangement. These ecological variations partially depend on the functional morphology of their digging tools. Muscular and mechanical features (e.g., lever arms relationship) of the bite force were analyzed in three caviomorph rodents with similar body size but different habits and ecological demands of the jaws. In vivo forces were measured at incisors' tip using a strain gauge load cell force transducer whereas theoretical maximal performance values, mechanical advantages, and particular contribution of each adductor muscle were estimated from dissections in specimens of Ctenomys australis (subterranean, solitary), Octodon degus (semi-fossorial, social), and Chinchilla laniger (ground-dweller, colonial). Our results showed that C. australis bites stronger than expected given its small size and C. laniger exhibited the opposite outcome, while O. degus is close to the expected value based on mammalian bite force versus body mass regressions; what might be associated to the chisel-tooth digging behavior and social interactions. Our key finding was that no matter how diverse these rodents' skulls were, no difference was found in the mechanical advantage of the main adductor muscles. Therefore, interspecific differences in the bite force might be primarily due to differences in the muscular development and force, as shown for the subterranean, solitary and territorial C. australis versus the more gracile, ground-dweller, and colonial C. laniger.

Evaluation of neophobia and its potential impact upon predator control techniques: a study on two sympatric foxes in southern Patagonia.

An alternative approach to increase the efficiency of predator control and selectivity is to consider the natural behavioural repertoire of the target species and how such behaviours may increase their vulnerability. Neophobia, or the hesitancy to approach a novel food item, object, or place, is an important factor influencing the investigative behaviour of animals, and its incorporation to predator control techniques may help to reduce losses of livestock to predators. In this study, we simultaneously evaluated the existence and intensity of neophobic responses in two sympatric fox species, the Culpeo (Pseudalopex culpaeus) and the Grey (P. griseus) foxes in southern Patagonia, Argentina. For this purpose, we used bait stations to compare fox behavioural responses in the absence (pre-treatment), presence (treatment) and removal (post-treatment) of a novel stimulus, which consisted of an orange PVC-traffic cone. Both fox species showed a neophobic response: bait-station visitation rates decreased (P=0.005 and P=0.048, for Culpeo and Grey foxes, respectively) in the presence of the novel object. The intensity of the response differed between species being higher for Culpeo foxes (approximately 80% of reduction in visitation rate during treatment for Culpeo foxes vs. 10% for Grey foxes). However, the bait-station visitation pattern after novel object removal indicated that animals probably increased exploration of the station. The high level of neophobia achieved by the Culpeo fox, together with an increase in post-treatment site exploration, suggests that behavioural manipulations (reduction of neophobia and its consequent increase in risk taking) could improve selective and efficient fox control in rural areas where livestock production is a major economic activity.

Biting performance and skull biomechanics of a chisel tooth digging rodent (Ctenomys tuconax; Caviomorpha; Octodontoidea).

Biting performance is a key factor in vertebrate groups possessing particular food habits. In subterranean rodents that use the incisors as a digging tool, apart from requirements related to gnawing abrasive diets, the force exerted at the incisors tips must be sufficient to break down soils that are often exceedingly compact. The subterranean genus Ctenomys diversified in the southern portion of South America closely associated with the relatively open environments that characterize that region. This genus is considered a "claw and chisel tooth digger," that is, during the excavation of their galleries, the animals break down the soil with both the fore-claws and the incisors. We report here measurements of in vivo bite force in one of the largest species of the genus, C. tuconax, which occupies highland grasslands with compacted soils. We document the combined use of claws and incisors observed under field conditions, also providing measurements of soil compaction in the habitat occupied by this species. We report estimates of bite force at the level of the incisors and cheek teeth calculated from the physiological cross-sectional area of jaw muscles. To this aim, anatomical and biomechanical analyses of the mandibular apparatus were performed in preserved specimens. We found that C. tuconax bites with a higher force than expected for a mammal of its size. To assess anatomical correlates of biting performance, the morphology of the skull and jaw, and incisor second moment of area were compared with those of other caviomorph rodents with different lifestyle.

Sexual selection in a polygynous rodent (Ctenomys talarum): an analysis of fighting capacity.

The South American subterranean rodent genus Ctenomys (Caviomorpha: Octodontoidea), which uses both claws and teeth to dig, shows striking morphological adaptations to its specialized mode of life. Among other traits, the genus has evolved a powerful jaw musculature and procumbent incisors that are used for dento-excavation. Behavioral observations indicate that these traits are also used during male aggressive encounters, which characterize the polygynous mating system of one of the species of the genus, Ctenomys talarum. A question emerges about sexual selection: could it have induced further changes in traits primarily evolved as adaptations for digging? To address this issue, we studied functional and morphological attributes of the jaw and incisors in specimens of C. talarum. Incisor bite forces were measured on wild females and males from a local population (Mar de Cobo; Buenos Aires Province) by means of a strain gauge load cell force transducer. Museum specimens coming from the same population were studied to assess anatomical attributes of both sexes. Since this species exhibits dimorphism in body size, the possible effect of body mass on the studied traits was analyzed. Males and females showed significant differences in biting performance and mandibular width, but when size was taken into account these differences disappeared. However, other dimorphic traits can vary with a certain independence with respect to size, particularly the 2nd moment of area of the incisors and, to a lesser extent, incisor procumbency. The former geometrical parameter, which is proportional to the bending strength, was highly dimorphic. This fact suggests that, during aggressive encounters between males, biting would place large bending loads on the incisors.

Scaling and adaptations of incisors and cheek teeth in caviomorph rodents (Rodentia, Hystricognathi).

The South American hystricognath rodents are one of the most diverse mammalian clades considering their occupied habitats, locomotor modes and body sizes. This might have been partly evolved by diversification of their masticatory apparatus' structure and its ecological commitment, for example, chisel-tooth digging. In this phylogeny-based comparative study, we test the relationship between ecological behavior and mechanical features of their incisors and molariforms. In 33 species of nine families of caviomorph rodents, we analyze incisor attributes related to structural stress resistance and molar features related with grinding capacity, for example, second moment of inertia and enamel index (EI) (enamel band length/occlusal surface area), respectively. Most of these variables scaled isometrically to body mass, with a strong phylogenetic effect. A principal component analysis discrimination on the EI clustered the species according to their geographic distribution. We presume that selective pressures in Andean-Patagonian regions, on particular feeding habits and chisel-tooth digging behaviors, have modeled the morphological characteristics of the teeth. Subterranean/burrower ctenomyids, coruros, and plains viscachas showed the highest bending/torsion strength and anchorage values for incisors; a simplified enamel pattern in molariforms would be associated with a better grinding of the more abrasive vegetation present in more open and drier biomes.

Inter-individual and age differences in exploration, neophobia and problem-solving ability in a Neotropical raptor (Milvago chimango).

Animal innovations have far-reaching ecological and evolutionary consequences. The occurrence and persistence of an innovation require several processes, including exploration, social and asocial learning, and low neophobia. In addition, the identity of the innovator may determine how these new behaviours are socially transmitted. Taking into account inter-individual and age differences, we investigated three correlates of animal innovation: object exploration, neophobia level and novel problem-solving ability in an opportunistic generalist raptor, the Chimango Caracara (Milvago chimango). Eighteen individuals (7 adults and 11 juveniles) were caught during the non-breeding period and housed in individual cages in outdoor aviaries. Each bird was given three tests: exploration, neophobia and problem-solving. Individuals differed in their response to novel situations both within and between age groups. Most of the juveniles were more explorative and had a lower neophobic response to a strange object than adult birds, but both age groups were able to solve a novel problem when given a food reward. In juveniles, neophobia level and problem-solving performance were inversely related; however, we found no relationship between these behaviours in adults. Exploration did not correlate with neophobia or problem-solving ability for either age group. This research is one of the few studies exploring the inter-individual and age differences in behavioural innovation and their correlates in a bird of prey. The explorative tendency, low neophobia and ability to innovate showed by M. chimango may be advantageous for this generalist and opportunistic raptor and might be some of the factors underlying its ecological success.

Evolution of brain size in a highly diversifying lineage of subterranean rodent genus Ctenomys (Caviomorpha: Ctenomyidae).

We evaluated brain size evolution in a specialized subterranean mammal, the ctenomyid rodent genus Ctenomys (tuco-tuco) and compared it, within a phylogenetic framework, to other caviomorph rodents differing in body size and modes of life. Although brain size in Ctenomys falls below the regression line obtained for caviomorph rodents, some fossorial species within the sister family Octodontidae, which make use of both above and below ground habitats, have a relatively larger brain size. Ctenomys has experienced a relatively recent and 'explosive' cladogenesis which yielded approximately 60 living species differing almost one order of magnitude in body size. Based on the evolutionary lag hypothesis between body versus brain size evolution, a negative correlation between encephalization quotient and body size would be expected for a group experiencing such a recent diversification in size. This is because a decrease of body size occurring during the emergence of a new species should produce a correspondingly greater encephalization and vice versa. Despite the fact that the allometric coefficient of brain weight versus body weight for 30 living species of Ctenomys was lower than that obtained for families within Caviomorpha, we failed to find any significant relationship between encephalization and body weight. Finally, the relationship between brain size and metabolism was assessed for those species of Ctenomys in which metabolic data are available. Brain size evolution in Ctenomys is discussed in regards to sensory capabilities and behavioral attributes associated with the occupation of a subterranean ecological niche.

Phylogeographical structure in the subterranean tuco-tuco Ctenomys talarum (Rodentia: Ctenomyidae): contrasting the demographic consequences of regional and habitat-specific histories.

In this work we examined the phylogeography of the South American subterranean herbivorous rodent Ctenomys talarum (Talas tuco-tuco) using mitochondrial DNA (mtDNA) control region (D-loop) sequences, and we assessed the geographical genetic structure of this species in comparison with that of subterranean Ctenomys australis, which we have shown previously to be parapatric to C. talarum and to also live in a coastal sand dune habitat. A significant apportionment of the genetic variance among regional groups indicated that putative geographical barriers, such as rivers, substantially affected the pattern of genetic structure in C. talarum. Furthermore, genetic differentiation is consistent with a simple model of isolation by distance, possibly evidencing equilibrium between gene flow and local genetic drift. In contrast, C. australis showed limited hierarchical partitioning of genetic variation and departed from an isolation-by-distance pattern. Mismatch distributions and tests of neutrality suggest contrasting histories of these two species: C. talarum appears to be characterized by demographic stability and no significant departures from neutrality, whereas C. australis has undergone a recent demographic expansion and/or departures from strict neutrality in its mtDNA.