Dental evolution in Neophanomys (Rodentia, Octodontidae) from the late Miocene of central Argentina

The evolutionary pattern of the molar morphology of the small caviomorph (Octodontidae) Neophanomys from the late Miocene Cerro Azul Formation of central Argentina is analyzed. Two new species (chronomorphs) are recognized, which constitute an anagenetically evolving lineage with a gradual and directional pattern of increasing molar hypsodonty. Dental changes related to increasing hypsodonty are comparable to those of the octodontid lineage Chasichimys also recovered from the Cerro Azul Formation. However, Neophanomys shows comparatively less variation in gross morphology and there are no evidences that this lineage achieved euhypsodonty. In contrast, important changes in enamel microstructure (schmelzmuster) are observed among different populations of Neophanomys, supporting the hypothesis that these changes can occur at least partially independently from modifications in dental gross morphology. The patterns of dental evolution detected in the Neophanomys and Chasichimys-Xenodontomys lineages and the unequivocal polarity of the changes involved, related to increasing hypsodonty, reinforce the hypothesis that chronological differences exist among late Miocene outcroppings of Cerro Azul Formation in central Argentina.     

Phylogenetic position ofAbalosia and the evolution of the extant Octodontinae (Rodentia, Caviomorpha, Octodontidae)

South American octodontid rodents of the subfamily Octodontinae currently show low species richness but great morphological and chromosomal diversity. This diversity is interpreted alternatively as the remnant of a wider past radiation or as the result of saltational evolution. These hypotheses are discussed in relation to a phylogenetic analysis of the Late Pliocene octodontineAbalosia castellanosi. My results suggest thatAbalosia, together withTympanoctomys andOctomys, is part of a clade of desert specialist propalinal octodontids, which would have differentiated east of the Andes in the emergent semi-deserts of western Argentina. The presence ofAbalosia in the coastal region of central Argentina during the Upper Marplatan Age (Late Pliocene) suggests a pulse of expansion of such arid environments, probably coeval with the global climatic deterioration detected around the transition Gauss-Matuyama magnetic ages. The phylogenetic position ofA. castellanosi suggests that extinction events affected the clade of the octodontine desert specialists. Accordingly, regardless of how rough or gradual the differentiation of the octodontine’s diversity has been, the living representatives seem to be remnants of a wider radiation.     

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.     

Arboreal gaits in three sympatric rodents Apodemus agrarius,Apodemus flavicollis (Rodentia,Muridae) and Myodes glareolus (Rodentia,Cricetidae)

Vertical stratification of the arboreal habitat allows the coexistence of several species in a given area, because the complex arboreal strata can be used in different ways by arboreal and scansorial mammals. The present report experimentally investigated the gait metrics on different arboreal substrates, of three sympatric rodents living in a deciduous forest in Poznań, Poland. Arboreal locomotion was compared between the burrowing striped field mouse, Apodemus agrarius, the scansorial bank vole, Myodes glareolus, and the more arboreal yellow-necked mouse, Apodemus flavicollis. We filmed two wild-caught individuals from each species walking on four different substrate diameters (2 mm, 5 mm, 10 mm, 25 mm) and three different inclinations (45° descending, horizontal, 45° ascending) at 240 fps and collected a set of gait parameters from a total of 273 complete cycles. Our results did not demonstrate clear relationships between arboreal locomotion and the ecology of the three species. Only A. flavicollis exhibited locomotor features partly associated with arboreal competence, including lower velocity and diagonality on narrow substrates and asymmetrical gaits on wider ones. On the other hand, the two Apodemus species, despite their different ecologies, shared a few locomotor similarities, such as velocity regulation primarily by stride frequency, and similar effects of substrate size and inclination on diagonality, duty factor, and duty factor index indicating the possibility of a phylogenetic signal. Because the selected gait parameters provided limited insight into the ability of small mammals to move competently through an arboreal habitat, these findings indicate that the relationship between behaviour and ecology is complex.     

Tooth morphology of Echimyidae (Rodentia,Caviomorpha): homology assessments,fossils, and evolution

Echimyidae constitute the most important radiation of caviomorph rodents in the Neotropical region, represented by 20 extant genera and several extinct species. Both in extant and fossil forms, this diversity is reflected by a significant morphological variation found in crown structures of the cheek teeth. Different hypotheses of primary homology have been proposed for these structures, which, in turn, support diverse dental evolutionary hypotheses. In this contribution we inspect the main structures (cusps and lophids) of the lower deciduous teeth and molars in extinct and extant Echimyidae, and establish their topological correspondences. Comparisons with cusps and lophids of Erethizontidae are emphasized. We explore the testing of alternative primary hypotheses of lophid correspondences in a cladistic context. Following a ‘dynamic’ approach, we select the hypothesis of primary homology, which produced the more parsimonious results, and evaluate the evolutionary transformations of the dental characters analysed. In this context, the phylogenetic relationships of living Myocastor coypus (Molina, 1782) with the extinct Tramyocastor and Paramyocastor are tested. Our results indicate that pentalophodonty is the derived condition for the lower molars in Echimyidae, that trilophodonty evolved independently at least three times during the evolutionary history of these rodents, and that tetralophodonty represents the plesiomorphic condition. This study shows that dental evolution in echimyids can be better understood when occlusal structures are expressed as reliably comparable characters, and when fossils are taken into account. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164 , 451–480.     

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.     

First complete skull of an octodontoid (Rodentia, Caviomorpha) from the Early Miocene of South America and its bearing in the early evolution of Octodontoidea

Octodontoidea is the most diverse group of caviomorph rodents. The systematics of most of the fossil representatives has been essentially based upon dental characters. Described here is an almost complete skull with dentition assigned to Prospaniomys Ameghino based upon its dental morphology. The specimen comes from the Sarmiento Formation at Pampa de Gan Gan (central Patagonia, Argentina), assigned to the Colhuehuapian SALMA (early Miocene). The most remarkable features are in the posterior portion of the skull, some of them shared with the modern octodontids and interpreted as specialized by previous authors, which contrast with the generalized dental morphology. These combined features were not previously known in other octodontoids. The comparisons with other fossil and extant members of the superfamily suggest that the characters traditionally used to associate Prospaniomys with the echimyids are very probably plesiomorphies. Prospaniomys would represent an early diverging lineage more closely related to modern octodontids than to echimyids, in which cranial structures evolved more rapidly than dental and mandibular ones.     

Phylogeny and adaptive diversity of rodents of the family Ctenomyidae (Caviomorpha): delimiting lineages and genera in the fossil record

Differentiation of genera of the modern (Late Miocene to Recent) South American rodent family Ctenomyidae would have been linked to the acquisition of disparate adaptations to digging and life underground. In accordance with this hypothesis, the delimitation of lineages and genera in the ctenomyid fossil record is evaluated here following an adaptation-rooted criterion that involves both an assessment of the monophyly and of the adaptive profiles of recognized clades. The application of such a criterion, including morphofunctional information, delimited four cohesive lineages among crown ctenomyids (i.e. euhypsodont species of the Late Miocene to Recent): Eucelophorus (Early Pliocene–Middle Pleistocene), Xenodontomys-Actenomys (Late Miocene–Pliocene), Praectenomys (Pliocene) and Ctenomys (including Paractenomys ; Pliocene–Recent); in addition, the results supported the status of Xenodontomys as a paraphyletic ancestor of Actenomys . The cladogenesis that gave rise to the crown group would have occurred immediately after the acquisition of euhypsodonty in a Xenodontomys simpsoni -like ancestor during the Late Miocene. This putative ancestor would have had fossorial habits and moderate digging specializations, an adaptive profile maintained in Xenodontomys-Actenomys . Eucelophorus and Ctenomys would have independently evolved subterranean habits at least since the Pliocene. Although the earliest history of the only living representative, Ctenomys , is known only fragmentarily, remains from Esquina Blanca (Uquía Formation), in north-western Argentina, suggest a minimum age of around 3.5 Ma (Early–Late Pliocene) for the differentiation of the genus. This date agrees with recent molecular estimates.     

Plio-Pleistocene evolution of the genus Castor (Rodentia, Mammalia) in Europe: C. fiber plicidens of Pietrafitta (Perugia, Central Italy)

In the lignite sediments of Pietrafitta (Tiberino Basin, Umbria, Central Italy), a rich fossil assemblage of vertebrate, invertebrate and plant remains belonging to the Farneta Faunal Unit (Late Villafranchian, Early Pleistocene) was collected. Among them fossil beaver cranial remains are relatively abundant. Western-Central Europe Villafranchian beaver remains were assigned to C. plicidens by some authors because molar occlusal surface patterns show complex enamel crenulations. Several recent authors have classified them as C. fiber while analysing other morphological patterns. Our samples have been compared to Plio-Pleistocene fossil remains and to living European populations of the genus Castor. New morphometric parameters on molar occlusal surfaces have been defined and statistical analyses (One-Way MANOVA, Principal Component Analysis and Canonical Discriminant Analysis) have been performed on them. The results point out a subspecific separation between the Late Villafranchian beaver of Pietrafitta, Quarata and San Giovanni in Valdarno (Umbria and Tuscany) and C. fiber populations. St. Vallier (France) Late Pliocene and Mosbach 2 (Germany) Middle Pleistocene beavers classified respectively by Viret and Friant as C. plicidens, show a C. f. fiber molar teeth pattern. Therefore, C. fiber plicidens did not occur in Central-Western Europe and this subspecific name may be used only for the local populations of Valdarno and Tiberino Basin (San Giovanni in Valdarno, Quarata, Pietrafitta and a few localities of the same area), at that time peripheral populations, probably semi-isolated during the Late Villafranchian, and at the southern limit of the geographic range of C. fiber.     

Functional morphology and paleobiology of the pliocene rodent Actenomys (Caviomorpha: Octodontidae): the evolution to a subterranean mode of life

The Pliocene caviomorph rodent Actenomys has long been recognized as an early fossorial representative of the subfamily Ctenomyinae (Octodontidae), which includes one living genus, Ctenomys (tuco-tucos), and several species widely distributed in South America. To assess the de gree of specialization for digging in Actenomys , we performed morphological comparisons with other octodontid genera (Octodon, Spalacopus , and Ctenomys) of known mode of life and behaviour. As a whole, our results indicate that, in terms of morphological specializations for digging, Actenomys occupies an intermediate position between Octodon , a generalized semi-fossorial rodent which forages above ground, and Ctenomys-Spalacopus , two highly specialized subterranean forms. The position of the deltoid process (humerus) and the length of the olecranon process (ulna)–two traits which affect the out-forces exerted by several forelimb muscles–were in Actenomys intermediate between non-subterranean and subterranean taxa. The skull, particularly the rostrum, appears to be strong, and sagittal and nuchal crests are well marked. The zygomatic arches arc as flared as those of the generalized Octodon. Notably, the upper and lower incisors of Actenomys were extremely procumbent, as in many highly-specialized chisel-tooth digging species. Based upon the stratigraphic and sedimentological analysis of the palaeosoils containing its fossil remains, we suggest that Actenomys lived in an environment of moderate to hig primary productivity. The texture of the palaeosoils indicate that theywere hard and highly cohesive. This situation, which contrasts with that observed in living Ctenomyinae, has relevant implications for burrowing cost. The integration of morphological and palaeoenvironmental data allowed testing of hypotheses about the palaeobiological attributes of this ancestral ctenomyine.     

Ancylotherium pentelicum (Perissodactyla, Mammalia) from the Upper Miocene of central and western Turkey

We report here new fossils of A. pentelicum from two Turolian localities of central and western Turkey, Pinaryaka and Salihpasalar (Kemikalan). The material, which consists of well preserved distal limb-bones, is compared with the few previously described specimens, mainly from Samos and Pikermi, but the observed differences should probably be attributed to individual variation. There is no evidence of evolution of the species during the Middle Turolian of the eastern Mediterranean, where it filled a narrow ecological niche, before its extinction.     

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. J. Morphol. 2012. © 2012 Wiley Periodicals, Inc.     

Unique differentiation of radial enamel in Arsinoitherium (Embrithopoda,Tethytheria)

The enamel microstructure in molars of Arsinoitherium is reinvestigated and a new modification of radial enamel (RE), ‘arsinoitheriid radial enamel (ARE)’, is defined. It is characterised by alternating stripes with different organisation of the interprismatic matrix but no prism decussation. Recognition of this new subtype leads to a reinterpretation of structure previously identified as modified radial enamel and of Hunter–Schreger bands in Arsinoitherium. The newly differentiated ARE of Arsinoitherium is more derived in relation to corresponding microstructures of Palaeoamasia and Crivadiatherium. A careful reinvestigation of RE in other Paenungulata will be required to provide additional data bearing on phylogenetic reconstruction. The enamel of Phenacolophus argues against inclusion of this genus in the Embrithopoda.     

Comparative study on the forefoot and hindfoot intrinsic muscles of some cavioidea rodents (Mammalia, Rodentia)

The present study compares the forefoot and hindfoot musculature of five representative species of Cavioidea rodents. In all species, the musculature of both forefeet and hindfeet have the same array regardless of the absence of digit I in the manus of Hydrochaeris hydrochaeris and Cavia porcellus. Our results suggest a tendency in these species towards a three-digit system, with a functional loss of digit V and a predominance of digit III in their forefeet. In the same way, the muscular reduction of digit I in the other rodents analyzed indicates a four-digit system with predominance of digit II in Myoprocta acouchy and Dasyprocta leporina and of digit V in Agouti paca. There seems to be an association between the muscular arrangement and functional axis of the foot, raising the general question why this axis runs between the third and forth digit, or along the third digit.     

Hypsosteiromys (Rodentia, Hystricognathi) from the Early Miocene of Patagonia (Argentina), the only Erethizontidae with a tendency to hypsodonty

Hypsosteiromys is the only New World porcupine that shows a tendency to hypsodonty. It is recorded exclusively from the Colhuehuapian Age (Early Miocene) of central Patagonia (Argentina). In addition to the type species, a second one, Hypsosteiromys nectus(AMEGHINO, 1902), is recognized, from the southern cliff of Colhuehuapi Lake (Chubut province). It differs from the type species in the lesser development of the anterolabial and posteroflexid notches of the m1-3, shorter p4 and dp4, and more slender incisors. Dental morphology suggests that the species of Hypsosteiromys lived in more open areas than most fossil and living Erethizontidae.     

Aeretesmelanopterus (Pteromyinae, Rodentia) from Tianyuan Cave near Zhoukoudian (Choukoutien) in China

The materials of Aeretesmelanopterus from the Tianyuan Cave, a Late Pleistocene site with human fossils recovered recently near Zhoukoudian, are the richest of this species ever reported in North China. Aeretesmelanopterus is an endemic species of China, its relationship with Petaurista and Trogopterus is still controversal. Studies based on the dental characters show that Aeretesmelanopterus resembles Petaurista more closely than Trogopterus. The earliest fossil record of Aeretesmelanopterus was found in South China in Middle Pleistocene deposits. The earliest records in the Beijing area are from Upper Cave and Tianyuan Cave at Zhoukoudian are of late-Late Pleistocene age, around 30 kyrs BP. The geographical distributions of this species are very limited. In evolution, Aeretesmelanopterus experienced both the increase and the decrease in tooth size; but its crown height of cheek teeth was continuously increasing.     

A new stage in the evolution of the mole rats (Rodentia,Spalacinae) from the Early Miocene of northern Greece

The cheek teeth of Debruijnia kostakii n. sp. from the Early Miocene of Karydia are only slightly smaller than those of the largest extant species of the Spalacinae (Spalax giganteus), but show characteristics that are reminiscent of its Paleogene cricetid ancestor. The peculiar combination of large size and primitive dental morphology of this species bridges the gap between Debruijnia and Pliospalax and thereby supports the hypothesis that these genera form the core of the complex evolutionary history leading to extant Spalax.     

Dental remains of Lower Miocene small rodents from Napak (Uganda): Afrocricetodontidae,Myophiomyidae, Kenyamyidae and Sciuridae

The Early Miocene sites of Napak (Uganda), which have been prospected by the Uganda Palaeontology Expedition since 1985, have yielded abundant fossil remains, including a rich and diverse rodent fauna. Ever since the work of Lavocat (1973) on the East African Miocene rodents, which focused mainly on the Kenyan remains (but included some material from Napak discovered by Bishop during the 1950s and 1960s), few studies have been made on the Ugandan specimens. This lack provides motivation for further study of Napak micro-mammals, especially the abundant rodents, which were collected at five sites, Napak IV, V, XV, XXX and XXXI. The fossils described here are attributed to six species representing four families: Afrocricetodontidae (Afrocricetodon songhori, Protarsomys macinnesi), Myophiomyidae (Myophiomys arambourgi), Kenyamyidae (Simonimys genovefae, Kenyamys mariae) and Sciuridae (Vulcanisciurus africanus). Comparison of the dentition of extant and extinct taxa allows us to propose dietary hypotheses (granivory, herbivory and omnivory) for these rodents. In addition, the fossil rodents associated with other fauna from the same localities suggest a more-or-less dense forest environment with clearings, and the probable presence of a humid climate at the time of deposition.