Size, shape and structural versatility of the skull of the subterranean rodent Ctenomys (Rodentia, Caviomorpha): functional and morphological analysis

Morphological analysis of the skull of the subterranean rodent Ctenomys , a highly speciose genus which uses both claws and teeth when digging, shows that for a broad range of species size, scaling was associated with both variation and maintenance of shape. Our results show that the angle of incisor procumbency (AIP), a character largely viewed as an adaptation to digging with teeth, is highly variable. We found a non-significant relationship between AIP and basicranial axis (basioccipital + basisphenoid) length, a measure of overall skull size. Accordingly, both small and large Ctenomys species possess either high or low AIP. A significant relationship between AIP and diastema length, given the rostral allometry seen in Ctenomys , suggests that hypermorphosis to a certain extent influences AIP. However, the roots of the incisor are lateral to those of the cheek teeth and their position may thus shift freely. This observation supports the notion that skull structural design, and to a certain extent rostral allometry, underlies variation in AIP. On the other hand, the positive allometry of incisor width and thickness indicates that, in larger species, proportionately powerful incisors are able to resist greater bending forces. We found that the out-lever arm of the jaw adductor muscles scales with positive allometry against basicranial axis length. However, we found an isometric relationship between in- and out-lever arms. In this case, conservation of skull proportions, regardless of variation in size, is a feature possibly related to the maintenance of an effective tooth digging capability. Functional and ecological data are discussed when assessing the implications of size and shape variation in the skull of Ctenomys .  © 2003 The Linnean Society of London. Biological Journal of the Linnean Society , 2003, 78 , 85−96.     

Bite force and body mass of the fossil rodent Telicomys giganteus (Caviomorpha,Dinomyidae)

An exceptionally well-preserved skull of the Pliocene rodent Telicomys giganteus allowed the first estimation of body mass and analysis of the bite mechanics of this species of South American giant rodent. In this study, we reconstructed the main anatomical features of the skull of this Pliocene rodent and related them to the bite force at the incisors. The average of an estimation body mass gives 100 kg. We also estimated the bite force using three different techniques. Two methods suggest that bite forces at the incisors have a range of 500–1000 N. However, the incisors seem to be stronger than expected for this bite force, implying that the bite forces may have been greater than 2000 N. We consider the hypothesis of defense against predators or other agonistic behavior to explain our results.     

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. J. Morphol. 275:902–913, 2014. © 2014 Wiley Periodicals, Inc.     

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.     

The relationship between skull morphology, biting performance and foraging mode in Kalahari lacertid lizards

Lizards are a diverse clade in which one radiation consists entirely of sit-and-wait foragers and another consists of wide foragers. Lizards utilizing these two foraging modes are known to differ in diet, but little is known about how feeding morphology relates to diet and/or foraging mode. This study tested the hypothesis that skull morphology and biting performance are related to diet preference, and consequently, coevolve with foraging mode. Four species of lacertid lizard were studied because they vary in foraging mode, their phylogenetic relationships are known and they are well studied ecologically. Using an 'ecomorphological' approach, skull morphology and biting performance were quantified and mapped on to the phylogeny for the species. The results indicate that sit-and-wait species have shorter, wider skulls than the wide foraging species, and that all are significantly different in overall head shape. The sit-and-wait species had similar values for biting performance; however, clear phylogenetic patterns of covariation were not present between sit-and-wait and wide foraging species for either biting performance or skull morphology. Thus, skull morphology and performance have little influence on diet and foraging mode in these species. Instead it is likely that other factors such as seasonal prey availability and/or life history strategy shape foraging mode decisions.  © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 140 , 403–416.     

Functional anatomy of the limbs of erethizontidae (Rodentia, Caviomorpha): Indicators of locomotor behavior in Miocene porcupines

Functional analysis of the limb bones of the erethizontid Steiromys duplicatus, one of the most abundant Miocene porcupines from Patagonia, provides evidence to infer their locomotor behavior. Remains of the giant Neosteiromys pattoni (Late Miocene of Northeast Argentina) are also analyzed. Osteological and myological features of extant porcupines were evaluated and used as a model to interpret the functional significance of Miocene species' limbs. Several features in erethizontids are compatible with the ability to climb: the low humeral tuberosities indicate a mobile gleno-humeral joint; the prominent and distally extended deltopectoral crest indicates a powerful pectoral muscle, which is particularly active when climbing; the humero-ulnar and humero-radial joints are indicative of pronation-supination movements; the well-developed lateral epicondylar ridge and the medially protruding entepicondyle are in agreement with an important development of the brachioradialis, supinator, flexor digitorum profundus, and pronator teres muscles, acting in climbing and grasping functions; the mechanical advantage of the biceps brachii would be emphasized because of its distal attachment on the bicipital tuberosity. As with extant porcupines, in Miocene species, the large femoral head would have permitted a broad range of abduction of the femur, and the medially protruding lesser trochanter would have emphasized the abduction and outward rotation of the femur by the action of the ilio-psoas complex. In S. duplicatus, the shape of the hip, knee, and cruro-astragalar, calcaneo-astragalar, and astragalo-navicular joints would have allowed lateral and rotational movements, although probably to a lesser degree than in extant porcupines. Foot features of S. duplicatus (e.g., great medial sesamoid bone, medial astragalar head, complete hallux) indicate that this species would have had grasping ability, but would not have achieved the high degree of specialization of Coendou. Steiromys duplicatus would have been a semiarboreal dweller, resembling Erethizon dorsatum.     

Evolution of bite force in Darwin's finches: a key role for head width

Studies of Darwin's finches of the Galapagos Islands have provided pivotal insights into the interplay of ecological variation, natural selection, and morphological evolution. Here we document, across nine Darwin's finch species, correlations between morphological variation and bite force capacity. We find that bite force correlates strongly with beak depth and width but only weakly or not at all with beak length, a result that is consistent with prior demonstrations of natural selection on finch beak morphology. We also find that bite force is predicted even more strongly by head width, which exceeds all beak dimensions in predictive strength. To explain this result we suggest that head width determines the maximum size, and thus maximum force generation capacity of finch jaw adductor muscles. We suggest that head width is functionally relevant and may be a previously unrecognized locus of natural selection in these birds, because of its close relationship to bite force capacity.     

The comparative gastrointestinal morphology of Jaculus jaculus (Rodentia) and Paraechinus aethiopicus (Erinaceomorpha)

Jaculus jaculus (Lesser Egyptian jerboa) and Paraechinus aethiopicus (Desert hedgehog) are small mammals which thrive in desert conditions and are found, among others, in the Arabian Peninsula. Jaculus jaculus is omnivorous while P. aethiopicus is described as being insectivorous. The study aims to describe the gastrointestinal tract (GIT) morphology of these animals which differ in diet and phylogeny. The GITs of J. jaculus (n = 8) and P. aethiopicus (n = 7) were weighed, photographed, and the length, basal surface areas, and luminal surface areas of each of the anatomically distinct gastrointestinal segments were determined. The internal aspects of each area were examined and photographed while representative histological sections of each area were processed to wax and stained using haematoxylin and eosin. Both species had a simple unilocular stomach which was confirmed as wholly glandular on histology sections. Paraechinus aethiopicus had a relatively simple GIT which lacked a caecum. The caecum of J. jaculus was elongated, terminating in a narrow cecal appendix which contained lymphoid tissue on histological examination. The internal aspect of the proximal colon of J. jaculus revealed distinct V‐shaped folds. Stomach content analysis of J. jaculus revealed mostly plant and seed material and some insects, whereas P. aethiopicus samples showed plant material in addition to insects, indicating omnivorous feeding tendencies in areas where insects may be scarce. J. Morphol. 277:671–679, 2016. © 2016 Wiley Periodicals, Inc.     

Testing Magnetic Orientation in a Solitary Subterranean Rodent Ctenomys talarum (Rodentia: Octodontidae)

To test for the hypothesis that Ctenomys talarum can use the earth's magnetic field for spatial orientation, we carried out field and laboratory experiments to analyse if C. talarum burrows present any geomagnetic orientation in their natural habitat, if C. talarum show any spontaneous directional preference when starting to excavate their burrows and if this subterranean rodent is capable to use the earth's magnetic field to orient towards a goal in a complex maze. No correlation between the burrowing direction and the earth's magnetic field was found. We could not find any evidence for any spontaneous directional preference when starting to excavate the burrows in C. talarum. The change of the horizontal vector of the geomagnetic field did not affect the ability of this rodent to orient towards a goal in an artificial labyrinth. Explanations for these results and other possible mechanisms of orientation that could be used by C. talarum are discussed.     

Cranial functional morphology of fossil dogs and adaptation for durophagy in Borophagus and Epicyon (Carnivora,Mammalia)

Morphological specialization is a complex interplay of adaptation and constraint, as similarly specialized features often evolve convergently in unrelated species, indicating that there are universally adaptive aspects to these morphologies. The evolutionary history of carnivores offers outstanding examples of convergent specialization. Among larger predators, borophagine canids were highly abundant during the tertiary of North America and are regarded as the ecological vicars of Afro‐Eurasian hyenas. Borophaginae is an extinct group of 60+ species, the largest forms evolving robust skulls with prominently domed foreheads, short snouts, and hypertrophied fourth premolars. These specializations have been speculated to enhance bone cracking. To test the extent that the skulls of derived borophagines were adapted for producing large bite forces and withstanding the mechanical stresses associated with bone cracking relative to their nonrobust sister clades, we manipulated muscle forces in models of six canid skulls and analyzed their mechanical response using 3D finite element analysis. Performance measures of bite force production efficiency and deformation minimization showed that skulls of derived borophagines Borophagus secundus and Epicyon haydeni are particularly strong in the frontal region; maximum stresses are lower and more evenly distributed over the skull than in other canids. Frontal strength is potentially coupled with a temporalis‐driven bite to minimize cranial stress during biting in the two derived genera, as tensile stress incurred by contracting temporalis muscles is dissipated rostro‐ventrally across the forehead and face. Comparison of estimated masticatory muscle cross section areas suggests that the temporalis‐masseter ratio is not strongly associated with morphological adaptations for bone cracking in Borophagus and Epicyon; larger body size may explain relatively larger temporalis muscles in the latter. When compared with previous studies, the overall cranial mechanics of the derived borophagines is more similar to bone‐cracking hyaenids and percrocutids than to their canid relatives, indicating convergence in both morphological form and functional capability. J. Morphol., 2010. © 2010 Wiley‐Liss, Inc.     

The influence of implant number and attachment type on maximum bite force of mandibular overdentures: a retrospective study

doi: 10.1111/j.1741‐2358.2010.00421.x
The influence of implant number and attachment type on maximum bite force of mandibular overdentures: a retrospective study Aim: Assessment of the influence of such factors as attachment type, number of implants, gender and age of patients on maximum bite force (MBF). Material and methods: Sixty‐two edentulous patients (32 females, 30 males; aged 64.03 ± 10.07 years, range 42–90 years) with mandibular implant overdentures with various attachment types were included in the study and their MBFs were recorded. The results were evaluated statistically at a significance level of p < 0.05. Results: The MBF in male patients was found to be statistically significantly (p < 0.05) higher than in female patients. No statistically significant differences (p > 0.05) were found with respect to age groups, attachment types and number of implants supporting the mandibular overdentures. Conclusion: Within the limitations of this study, it is concluded that independent of the number applied, dental implants increase MBF in edentulous patients. While males show higher bite force, patient age and attachment type seem not to play an important role.     

Inter and intra-specific hybridization in tuco-tucos (Ctenomys) from Brazilian coastal plains (Rodentia: Ctenomyidae)

The present work describes chromosomal polymorphisms in zones of contact between divergent populations of Ctenomys minutus parapatrically distributed in the coastal plain of southern Brazil, and inter-specific hybridization with C. lami a closely related species. A sample of 171 specimens from 32 sample sites distributed along 161 km of the coastal plain was cytogenetically analyzed. Nine polymorphic populations were found: four with specimens with 2n = 46–48 (autosomal arm number (AN) = 76); three only have specimens with 2n = 47 and 48; one population sampled presented specimens with 2n = 43–46 (AN = 74–76) and one population with 2n = 50–52 (AN = 76–80). The remainder populations were fixed for 2n = 42, 46 or 48. The variation is the result of Robertsonian mechanisms of chromosomal evolution and a fusion in tandem rearrangement. The polymorphisms have been considered the result of secondary contact of populations after divergence in allopatry. The geomorphological evolution of the coastal plain provides clues to the possible existence of past geographic barriers acting over populations of Ctenomys, during the Holocene.     

Chromosome plasticity in Ctenomys (Rodentia Octodontidae): chromosome 1 evolution and heterochromatin variation

A chromosome 1 (Cr1) pericentric inversion is described in six of seven species in the genus Ctenomys (tuco-tucos) from Uruguay. The inversion was inferred from G-band analyses of subtelocentric Cr1 hypothesised to be derived from the ancestral metacentric condition. Cr1 varies across species in heterochromatin amount and localisation including a metacentric chromosome without positive C-bands in C. torquatus, a subtelocentric chromosome with heterochromatic short arms in C. rionegrensis, and a subtelocentric chromosome negative after C-banding in five of the species analysed here. Pachytene chromosomes from C. rionegrensis, a species with the highest heterochromatin content, and C. torquatus, one of the species with the lowest heterochromatin content, were analysed in order to assess possible mechanisms of heterochromatin evolution. This analysis revealed the presence of three heterochromatic chromocenters in C. rionegrensis where bivalents converge, while in C. torquatus only one chromocenter was observed. In both species, highly repetitive DNA was observed, localised in chromocenters after “in situ” hybridisation. Heterochromatin associated protein M31 was localised in chromocenters of both species after immuno-detection. The spread of heterochromatin in Ctenomys chromosomes could be produced by chromatin exchanges at the chromocenter level. We propose the exchange of this DNA associated proteins between non-homologous chromosomes in pachytene to be the responsible for the spread of heterochromatin through the karyotypes of species like C. rionegrensis     

Early evolutionary differentiation of morphological variation in the mandible of South American caviomorph rodents (Rodentia, Caviomorpha)

Caviomorphs are a clade of South American rodents recorded at least since the early Oligocene (> 31.5 Ma) that exhibit ample eco-morphological variation. It has been proposed that phylogenetic structure is more important than ecological factors for understanding mandibular shape variation in this clade. This was interpreted as a result of the long-standing evolutionary history of caviomorphs and the early divergence of major lineages. In this work, we test this hypothesis through the analysis of morphological variation in the mandible of living and extinct species and compare this information with that obtained through comparative phylogenetic analyses. Our results support the hypothesis of early origin of mandibular variation; moreover, they suggest the conservation of early differentiated morphologies, which could indicate the existence of constrained evolutionary diversification.     

Locomotory adaptations in entoptychine gophers (Rodentia: Geomyidae) and the mosaic evolution of fossoriality

Pocket gophers (family Geomyidae) are the dominant burrowing rodents in North America today. Their fossil record is also incredibly rich; in particular, entoptychine gophers, a diverse extinct subfamily of the Geomyidae, are known from countless teeth and jaws from Oligocene and Miocene-aged deposits of the western United States and Mexico. Their postcranial remains, however, are much rarer and little studied. Yet, they offer the opportunity to investigate the locomotion of fossil gophers, shed light on the evolution of fossoriality, and enable ecomorphological comparisons with contemporaneous rodents. We present herein a quantitative study of the cranial and postcranial remains of eight different species of entoptychine gophers as well as many contemporary rodent species. We find a range of burrowing capabilities within Entoptychinae, including semifossorial scratch-digging animals and fossorial taxa with cranial adaptations to burrowing. Our results suggest the repeated evolution of chisel-tooth digging across genera. Comparisons between entoptychine gophers and contemporaneous rodent taxa show little ecomorphological overlap and suggest that the succession of burrowing rodent taxa on the landscape may have had more to do with habitat partitioning than competition.     

Mandibular corpus strain in primates: Further evidence for a functional link between symphyseal fusion and jaw-adductor muscle force

Previous work indicates that compared to adult thick-tailed galagos, adult long-tailed macaques have much more bone strain on the balancing-side mandibular corpus during unilateral isometric molar biting (Hylander [1979a] J. Morphol. 159:253–296). Recently we have confirmed in these same two species the presence of similar differences in bone-strain patterns during forceful mastication. Moreover, we have also recorded mandibular bone strain patterns in adult owl monkeys, which are slightly smaller than the galago subjects. The owl monkey data indicate the presence of a strain pattern very similar to that recorded for macaques, and quite unlike that recorded for galagos. We interpret these bone-strain pattern differences to be importantly related to differences in balancing-side jaw-adductor muscle force recruitment patterns. That is, compared to galagos, macaques and owl monkeys recruit relatively more balancing-side jaw-adductor muscle force during forceful mastication. Unlike an earlier study (Hylander [1979b] J. Morphol. 160:223–240), we are unable to estimate the actual amount of working-side muscle force relative to balancing-side muscle force (i.e., the W/ B muscle force ratio) in these species because we have no reliable estimate of magnitude, direction, and precise location of the bite force during mastication. A comparison of the mastication data with the earlier data recorded during isometric molar biting, however, supports the hypothesis that the two anthropoids have a small W/ B jaw-adductor muscle force ratio in comparison to thick-tailed galagos. These data also support the hypothesis that increased recruitment of balancing-side jaw-adductor muscle force in anthropoids is functionally linked to the evolution of symphyseal fusion or strengthening. Moreover, these data refute the hypothesis that the recruitment pattern differences between macaques and thick-tailed galagos are due to allometric factors. Finally, although the evolution of symphyseal fusion in primates may be linked to increased stress associated with increased balancing-side muscle force, it is currently unclear as to whether the increased force is predominately vertically directed, transversely directed, or is a near equal combination of these two force components (cf. Ravosa and Hylander [1994] In Fleagle and Kay [eds.]: Anthropoid Origins. New York: Plenum, pp. 447–468). Am J Phys Anthropol 107:257-271, 1998. © 1998 Wiley-Liss, Inc.     

Musculoskeletal underpinnings to differences in killing behavior between North American accipiters (Falconiformes: Accipitridae) and falcons (Falconidae)

Accipiters (Accipiter spp.) and falcons (Falco spp.) both use their feet to seize prey, but falcons kill primarily with their beaks, whereas accipiters kill with their feet. This study examines the mechanistic basis to differences in their modes of dispatching prey, by focusing on the myology and biomechanics of the jaws, digits, and distal hindlimb. Bite, grip, and distal hindlimb flexion forces were estimated from measurements of physiological cross-sectional area (PCSA) and indices of mechanical advantage (MA) for the major jaw adductors, and digit and tarsometatarsal flexors. Estimated bite force, total jaw adductor PCSA, and jaw MA (averaged over adductors) tended to be relatively and absolutely greater in falcons, reflecting their emphasis on biting for dispatching their prey. Differences between genera in estimated grip force, total digit flexor PCSA, and digit MA (averaged over inter-phalangeal joints and digits) were not as clear-cut; each of these parameters scaled positively allometric in accipiters, which may reflect the scaling of both prey size, and the proportion of mammalian prey consumed by this lineage with increasing body size. Estimated tarsometatarsal force was greater in falcons than in accipiters, due to their greater MA, which may reflect selection for incurring greater forces during prey strikes. Conversely, the comparatively lower tarsometatarsal MA in accipiters reflects their capacity for greater foot speed potentially necessary for grasping elusive prey. Thus, this study elucidates how differences in jaw and hindlimb musculoskeletal morphology of accipiters and falcons are reflected in differences in their killing modes, and through differences in their force-generating capacities.